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SOUTH 

AUSTRALIAN 

MUSEUM 






CONTENTS PAGES 

BARTON, D. P. 13-30 

A checklist of helminth parasites of Australian Amphibia. 

BOLES, W. E. & MACKNESS, B. 139-149 

Birds from the Bluff Downs Local Fauna, Allingham Formation, Queensland, 

BURROW, C. 175-185 

Form and function in scales of Ligulalepis toombsi Schultze, an enigmatic palaeoniscoid 
from the early Devonian of Australia. 

CREASER, P. 215-216 

Protection of Movable Cultural Heritage 

JINMAN, M. 213-214 

World Heritage and fossils. 

LONG, J. & MACKNESS, B. 95-110 

Studies of the Late Cainozoic diprotodontid marsupials of Australia. 4. The Bacchus Marsh 
Diprotodons - geology, sedimentology and taphonomy. 

MCGOWRAN, B. & LI, Q. 197-212 

The Miocene oscillation in Southern Australia. 

MCNAMARA, G. 187-196 

Cape Hillsborough: an Eocene-Oligocene vertebrate fossil site from northeastern Queensland. 

MCNAMARA, J. A. 111-115 

A new fossil wallaby (Marsupialia: Macropodidae) from the southeast of South Australia. 

PARKER, S A. & COOK, P. L. 1-1 1 

Records of the bryozoan family Selenariidae from Western Australia and South Australia, with 
the description of a new specis of Selenaria Busk, 1854. 

PLEDGE, N. S. 65-77 

Fossils of the Lake: a history of Lake Callabonna excavations. 

PLEDGE, N. S. 117-123 

Cetacean fossils from the Lower Oligocene of South Australia. 

SUTTON, P. 31-52 

Material culture traditions of the Wik people, Cape York Peninsula. 

TEDFORD, R. H. 79-93 

Succession of Pliocene through medial Pleistocene mammal faunas of southeastern Australia. 

THULBORN, A. 151-158 

Mimicry in ankylosaurid dinosaurs. 

TYLER, M. J., GODTHELP, H. & ARCHER, M. 169-173 

Frogs from a Plio-Pleistocene site at Floraville Station, northwest Queensland. 

VANTETS, G. F. 135-138 

An extinct new species of cormorant (Phalacrocoracidae: Aves) from a Western Australian 
peat swamp. 

WHITE, A. W. & ARCHER, M. 159-167 

Elseya lava racko rum, a new Pleistocene turtle from fluviatile deposits at Riversleigh, 
north-western Queensland. 

WORTHY, T. H. 125-134 

Late Quaternary changes in the moa fauna (Aves: Dinornithiformes) on the West Coast of 
the South Island, New Zealand. 

ZBIK, M. & PRING, A. 53-56 

The Choolkooning 001 meteorite: a new (L6) olivine-hypersthene chondrite from South Australia. 

ZEIDLER, W. 57-64 

Obituary - Shane A. Parker. 

Volume 27(1) was published on 5 July 1994. 
Volume 27(2) was published on 28 October 1994. 

ISSN 0376-2750 



RECORDS OF THE BRYOZOAN FAMILY SELENARIIDAE FROM 

WESTERN AUSTRALIA AND SOUTH AUSTRALIA, WITH THE 

DESCRIPTION OF A NEW SPECIES OF SELENARIA BUSK, 1854 

S. A. Parker & P. L. Cook 



Summary 

Unlike that of Bass Strait, the selenariid of southern Western Australia and South Australia is 
relatively poorly known. A first estimate of its diversity is presented here, based on a study of 514 
colonies (in 100 samples), mostly previously unreported species collected by Sir Joseph Verco 
(1851 - 1935). The study confirms an earlier suspicion that Selenaria hexagonalis Maplestone, 1904 
actually consists of two species, here distinguished as S. hexagonalis sensu stricto and S. verconis 
sp. nov. In all, 16 species are represented, of which 11 now constitute new records for the region 
(South Australia: Otionella nitida (Maplestone, 1909) O. australis Cook & Chimonides, 1985b, 
Selenaria punctata Tenison-Woods, 1880, S. concinna Tenison-Woods, 1880, S. varians Cook & 
Chimonodes, 1987, S. exasperus Cook & Chimonides, 1987, S. verconis sp. nov. and 'S'. alata 
auctt. (non Tenison-Woods, 1880); Western Australia: S. bimorphocella Maplestone, 1904, S. 
concinna, S. hexagonalis and S. verconis sp. nov). Brief description of the genera and species are 
provided. 



RECORDS OF THE BRYOZOAN FAMILY SELENARIIDAE FROM WESTERN AUSTRALIA 
AND SOUTH AUSTRALIA, WITH THE DESCRIPTION OF A NEW SPECIES OF SELENARIA 

BUSK, 1854 



S. A. PARKER & P. L. COOK 



PARKER, S. A. & COOK, P. L. 1994. Records of the bryozoan family Selenariidae from Western 
Australia and South Australia, with the description of a new species of Selenaria Busk, 1854. Rec. S. 
Aust. Mus.21{\)\ 1-11. 

Unlike that of Bass Strait, the selenariid fauna of southern Western Australia and South Australia is 
relatively poorly known. A first estimate of its diversity is presented here, based on a study of 514 
colonies (in 100 samples), mostly previously unreported specimens collected by Sir Joseph Verco 
(1851-1935). The study confirms an earlier suspicion that Selenaria hexagonalis Maplestone, 1904 
actually consists of two species, here distinguished as S. hexagonalis sensu stricto and S. verconis sp. 
nov. In all, 16 species are represented, of which 11 constitute new records for the region (South 
Australia: Otionella nitida (Maplestone, 1909), O. australis Cook & Chimonides, 1985b, Selenaria 
punctata Tenison-Woods, 1880, S. concinna Tenison- Woods, 1880, S. varians Cook & Chimonides, 
1987, S. exasperans Cook & Chimonides, 1987, S. verconis sp. nov. and '5". alata auctt. (non 
Tenison-Woods, 1880); Western Australia: S. bimorphocella Maplestone, 1904, S. concinna, S. 
hexagonalis and S. verconis sp. nov). Brief descriptions of the genera and species are provided. 

S. A. Parker 1 , South Australian Museum, North Terrace, Adelaide, South Australia 5000, and P. L. Cook, 
Associate, Museum of Victoria, Swanston Street, Melbourne, Victoria 3000. Manuscript received 22 
March 1993. 



The calcareous element of southern Australian 
shelf sediments contains in places a high proportion 
of living bryozoans and dead bryozoan skeletons 
(Wass et al. 1970, James et at. 1992). Although 
many bryozoans live in or on such sediments, the 
free-living cup-shaped (lunulitiform) species of the 
anascan family Selenariidae can be particularly 
abundant in this zone. However, no idea of the 
abundance and diversity of selenariids in southern 
Australia was received until the appearance of Cook 
and Chimonides's studies on the Australasian 
Selenariidae (Cook & Chimonides 1984a, b, 1985a- 
c, 1986, 1987). From the Tertiary to the Recent of 
southern Australia (mainly Victoria and Bass 
Strait), these authors listed four genera (one new) 
and 43 species (18 new), many of the latter with an 
extensive temporal, and sometimes a wide 
geographical, range. 

The marine invertebrate collections of the South 
Australian Museum include much material obtained 
by Dr Sir Joseph Verco in the years 1890-1912. 
Verco was principally interested in molluscs, and 
the chief method he employed to obtain these 
(dredging) resulted in the collection of other benthic 
groups such as brachiopods (Verco & Blochmann 



1 (Shane Parker died in November 1992. This 
paper was then in the final stages of preparation, 



P.L.C. 



1910), turbinoliid scleractinians (Cairns & Parker 
1992) and selenariid bryozoans. 

In 1904, in correspondence to Sir Sidney Harmer, 
C. M. Maplestone mentioned Verco's selenariid 
collections, and offered to have specimens of his 
own new species Selenaria hexagonalis and 5. 
bimorphocella, sent to Harmer by Verco. 
Subsequently, Livingstone (1928), in discussing 
Verco's bryozoans in the SAM, listed two samples 
under Selenaria punctata Tenison-Woods and one 
under Lunularia capulus (Busk); comments on these 
identifications are given below. 

Apart from these instances, Verco's selenariids 
have remained unreported. Inasmuch as his 
collecting stations ranged from the south-east of 
South Australia (off Beachport and Cape Jaffa) 
westward to the Albany and King George Sound 
districts of Western Australia, they complement 
those reported by Cook and Chimonides, which 
were mainly in the Bass Strait, with one off Jurien 
Bay near Perth. 

Verco's selenariid material in the SAM numbers 
48 samples. This was augmented by other material, 
including 19 Verco samples in the QM, four 
samples from off Perth, 1963 (QM), four from 
Western Australia (WAM) and four from South 
Australia, 1982-1991 (SAM). The following article 
lists the 16 species involved, together with 
synonymies, references to recent redescriptions, 
details of material examined, notes on Recent and 



S. A. PARKER & P. L.COOK 



fossil distribution and bathymetric range, and 
remarks on the distinguishing features of the species 

and genera. 



Materials and methods 

Abbreviations of institutions referred to in this 
paper are: BMNH, Natural History Museum, 
London; NMV, Museum of Victoria, Melbourne; 
QM, Queensland Museum, Brisbane; SAM, South 
Australian Museum, Adelaide; WAM, Western 
Australian Museum, Perth, 

Five hundred and fourteen specimens (colonies) 
in 100 samples were examined, of which 406 
specimens in 69 samples were collected by Sir 
Joseph Verco. By institution, the material was 
constituted as follows: SAM: Verco Coll. 268(48), 
other sources 78(20); QM: Verco Coll. 136(19), 
other sources 5(4); NMV: Verco Coll. 1(1), other 
sources 1(1); BMNH: Verco Coll. 1(1), other 
sources 2(2); WAM: other sources 22(4). Among 
the specimens examined were five syntypes of 
Lunulites patelliformis Maplestone, 1904 (= 
Lunularia capulus Busk, 1852a) and the lectotype 
of Selenaria hexagonalis Maplestone, 1904. All 
specimens were Recent except for a Pliocene 
paratype of S. verconis. Unless otherwise specified, 
all specimens referred to under Material Examined 
were collected by Verco. 



affinis thrives at nearly 250 m off New Zealand. 
Generally, the most abundant and diverse living 
fauna seems to occur between 50 and 130 m depth. 
The material examined here shows few exceptions, 
but nearly all the best preserved colonies with 
mandible and frontal membranes intact, which may 
be inferred to have been alive when collected, occur 
from depths shallower than 60 m. The exceptions 
are Helixotionella spiralis and H. scut at a, which 
together with S, pulchella were all originally 
collected alive from off Jurien Bay, Western 
Australia at 137 m. All three species were collected 
alive by Verco from off Albany at 147 m (see 
below), which suggests that their tolerances are 
normally at the deeper end of the range. 



Systematics 

Order Cheilostomatida Busk, 1 852b 
Suborder Anasca Levinsen, 1909 
Superfamily MICROPOROIDEA Gray, 1848 
Family SELENARIIDAE Busk, 1854 

Genus Lunularia Busk, 1 884 

Zooids large with limited cryptocyst, budded 
radially; avicularia large, simple, with paddle- 
shaped mandibles; large brooding zooids scattered. 



Notes on lunulitiform colonies 

Although some lunulitiform colonies are attached 
by rhizoids to shell fragments etc., on the sea- 
bottom, the free living species are unattached and 
are stabilized and supported by the elongated 
avicularian mandibles of the peripheral regions of 
the colony. Some species are capable of locomotion 
(see Cook & Chimonides, 1978 and Chimonides & 
Cook, 1981). They live in, or upon the upper 
centimetres of sediment, where their dead skeletons 
also accumulate. Colonies rarely exceed 25 mm in 
diameter and are conical, cup-shaped or discoid. 
Each colony, whether spirally or radially budded, 
has patterned groups of feeding zooids (autozooids) 
and avicularia, which have elongated, paddle-shaped 
or whip-like mandibles. Large brooding zooids tend 
to be found subperipherally or peripherally, and in 
the genus Selenaria, very large, specialized, non- 
feeding male zooids occur peripherally among 
enlarged avicularia. 

Some species, or populations within species, 
seem to have particular depth and/or temperature 
tolerances. For example, living Selenaria maculata 
is common in only 2—4 m in Queensland, but has 
been found at 146 m in New South Wales; Otionella 



Lunularia capulus (Busk, 1852a) 

Lunulites capulus Busk, 1852a: pi. 1, figs 13, 14. 
1854: 100, pi. 112. 

Lunulites gibbosa Busk, 1854: 100, pi. 111. 
Lunulites patelliformis Maplestone, 1904: 215, pi. 
25, fig. 6. 

Selenaria livingstonei Bretnall, 1922: 190, figs 2, 

2a. 

Lunularia capulus: Livingstone 1924: 198, 1928: 

1 15; Cook & Chimonides, 1986: 691, figs 6, 9, 12- 

14. 

Material Examined 

South Australia: Petrel Bay, St Francis I., 19 
fms (34.8 m), SAM L532(l); 13 Nm (nautical 
miles) (23.8 km) ESE of Troubridge Point, 35 m, K. 
L. Gowlett-Holmes and S. Corigliano 11- 
12.vi.1991, SAM L675(l); Investigator Strait, 20 
fms (36.6 m), SAM L395(l); Gulf St Vincent, SAM 
L492-496(5) (syntypes of Lunulites patelliformis 
Maplestone); W of Aldinga, Gulf St Vincent, 38-41 
m, K. L. Gowlett-Holmes and S. Corigliano 4- 
5.V.1987, SAM L533(l); Emu Bay, Kangaroo I., ca 
35 m, J. Gehling 4.iv.l984, sandy bottom, SAM 
L549(2); no data, SAM L535, 536(2). 



THE BRYOZOAN FAMILY SELENARIIDAE 



Distribution 

Previously known from Torres Straits, 
Queensland, New South Wales, Victoria, Bass Strait 
(including Bank Strait between Flinders I. and 
Tasmania), South Australia and south-western 
Western Australia, 27.5-167 m, with fossil records 
from the Miocene and Pliocene of Victoria and the 
Pliocene of South Australia and Western Australia. 
Previous South Australian records are from 
Investigator Strait, Gulf St Vincent and Backstairs 
Passage; the present material adds St Francis I. and 
Kangaroo I. to the known range in that State. 

Remarks 

Of the 13 colonies examined, seven were alive 
when collected. L. capulus is characterized by its 
large, deeply domed zoaria, and autozooids and 
avicularia alternating in distally contiguous radial 
series. The three colonies of L549(2) and L675(l) 
are the largest so far seen by us, measuring 34, 34 
and 35 mm in diameter respectively. 

One of the two colonies of L549 bears on its 
basal surface good examples of basal buds 
(structures previously described by Cook & 
Chimonides, 1986: 697-698). 

The sheltered, strongly concave basal surface of 
L. capulus is often colonized by other bryozoans. In 
the present specimens, the most frequent of these is 
the microporid Mollia multijuncta (Waters, 1879), 
which occurs in samples L492, L532, L549 and 
L675. The two zoaria of L549 bear also basal 
colonies of a species of Arachnopusia Jullien, 1888, 
Chorizopora brongniartii (Audouin, 1826), 
Microporella lunifera Haswell, 1880 and three 
species of Parasmittina Osburn, 1952. Apart from 
the small area of M. multijuncta, L675 is almost 
completely encrusted basally with Membranipora 
perfragilis (MacGillivray, 1881), and supports also 
a small colony of Scrupocellaria sp. and a minute 
individual (3 mm diameter) of a scleractinian coral, 
probably Scolymia australis (Milne Edwards & 
Haime, 1849). On the basal surface of L533 occurs 
yet another bryozoan, Crassimarginatella pyrula 
(Hintks, 1881). 

Cook (1985: 23-24, 93) discussed the occurrence 
of acrothoracid cirripedes in large lunulitiform 
bryozoans, particularly that of Kochlorinopsis 
discoporellae Stubbings, 1967 in West African 
populations of the cupuladriid bryozoan 
Discoporella umbellata (Defrance, 1823). In our 
material, these barnacles have been present in 
samples L492, L532, L533 and L675, as attested by 
the small slits at the apex of the colonies. 

Lunularia repanda (Maplestone, 1904) 

Lunulites repandus Maplestone, 1904: 216, pi. 25, 

fig- 7. 



Lunularia repandus: Cook & Chimonides, 1986: 
698, figs 7, 8, 10, 15, 16. 

Material Examined 

Western Australia: 32°S., l^OSE., off Perth, 
119 m, B. Jamieson 28.viii. 1963, QM GH1 150(1); 
King George Sound, 12-14 fms (22-25.6 m), SAM 
L537(l), 28 fms (51 m), SAM L538(l), 35 fms (64 
m), QM GH1693(1); 80 Nm (146.4 km) W of Eucla, 
81 fms (148 m), iii.1912, SAM L539(l), 140 fms 
(256 m), SAM L540(l). 

South Australia: Off St Francis 1., 35 fms (64 
m), SAM L541(l); cove NW of Petrel Cove, St 
Francis I., 40 m shifting sandy bottom, W. Zeidler 
& N. Holmes 28. i. 1982, SAM L548(3); S of 
Troubridge I., 20 fms (36.6 m), SAM L542(l); 
Beachport, 150 fms (275 m), SAM L543(l); no 
data, SAM L534(l), L544-547(36). 

Distribution 

Previously known from south-western Western 
Australia, South Australia and Bass Strait (27.5- 
183 m) and from the Kermadec Ridge (145-350 m; 
see Gordon, 1984), and by fossils from the Miocene 
of New Zealand. The present material adds King 
George Sound and 80 Nm (146.4 km) W of Eucla to 
the known Western Australian range, and St Francis 
I. and Beachport to the known South Australian 
range, and increases the recorded depth-range to 22- 
275 m. 

Remarks 

Of the 49 colonies examined, 29 were alive when 
collected. L. repanda differs from L. capulus in 
having flatter zoaria, and the autozooids in radial 
series not alternating with series of the avicularia 
which are very large and scattered. One sample 
(SAM L547) comprised 18 colonies, several of 
which exceeded 29 mm in diameter. Colonies in 
SAM L542 and L548 bore the bryozoan Mollia 
multijuncta (Waters, 1879) (Microporidae) on their 
basal surfaces. 

Genus Otionella Canu & Bassler, 1917 

Zooids budded radially, with small, rounded 
opesiae. Ancestrula with one distal and one 
proximal adjacent avicularium. Brooding zooids 
enlarged, marginal. 

Otionella squamosa (Tenison-Woods, 1880) 

Selenaria squamosa Tenison-Woods, 1880: 29, fig. 
29. 

Otionella squamosa: Cook & Chimonides, 1984a: 
232, figs 2, 6, 7, 14 g,h, 15, 21E; 1985b: 586, figs 
16, 17; Gordon, 1986: 71, pi. 28C. 



S. A. PARKER & P. L. COOK 



Material Examined 

?South Australia: no data, SAM L550(l). 

Distribution 

O. squamosa has been reported from Torres 
Strait, New South Wales, Bass Strait (20-121 m) 
and New Zealand (1092.5 m), and as a fossil from 
Victoria and Bass Strait (Pliocene) an^ New 
Zealand (Pleistocene). The present specimen, 
though without details of provenance, is most likely 
from South Australia. 

Remarks 

O. squamosa is distinguished by its large, 
scattered, usually asymmetrical avicularia, with 
marginally perforated cryptocyst, and prominent 
condyles unfused or fused on the basal side only. 

Otionella nitida (Maplestone, 1909) 

Selenaria nitida Maplestone, 1909: 271, pi. 77, fig. 

8. 

Otionella nitida: Cook & Chimonides, 1984a: 239, 

figs 14f, 16, 18-20, 21B; 1985b: 584, figs 14, 15, 

29. 

Material Examined 

South Australia: Backstairs Passage, 'deep 
water', SAM L551(l); Cape Jaffa, QM GH1642(1); 
no data, SAM L552(4), L553(l). 

Distribution 

Previously known from south-western and mid- 
western Western Australia, Bass Strait and New 
South Wales (17-148 m), with fossil records from 
south-western Western Australia (Pliocene) and 
New Zealand (Pleistocene, Miocene). The present 
material adds South Australia to the Recent 
distribution of the species. 

Remarks 

Of the seven colonies examined, two were alive 
when collected. Colonies of O. nitida are very small 
and domed; the small, symmetrical avicularia, 
which may occur in contiguous radial series, have a 
perforated cryptocyst and fused condyles. 

Otionella australis Cook & Chimonides, 1985b 

Otionella australis Cook & Chimonides, 1985b: 
590, figs 11, 12,23-25. 

Material Examined 

Western Australia: 32°S., 115°08'E., off Perth, 
119m, B. Jamieson 28.viii. 1963, QM GH1 150(2); 
King George Sound, 35 fms (64 m), QM 
GH 1693(3); 80 Nm (146.4 km) W of Eucla, 81 fms 
(148 m), in. 1912, SAM L554(49); W of Eucla, 50- 



120 fms (92-220 m), iii.1912, SAM L555(4); 40 
Nm (73.2 km) W of Eucla, 72 fms (132 m), iii.1912, 
SAM L556(2). 

South Australia: Cape Jaffa, QM GH 1642(55); 
off Cape Jaffa, 90 fms (165 m), SAM L557(l), 130 
fms (238 m), SAM L558(l); no data, SAM L559(l), 
L560(2). 

Distribution 

Previously known from south-western Western 
Australia and Bass Strait, 84-137 m and in the 
fossil record from south-western Western Australia 
(Pliocene) and Victoria (Miocene). The present 
material adds South Australia (south-eastern) and 
the Western Australian sector of the Great 
Australian Bight to the species' known distribution, 
and extends the depth-range to 238 m. 

Remarks 

Of the 62 colonies in the SAM, 25 were alive 
when collected. O. australis is distinguished by the 
simple, scattered, almost symmetrical avicularia 
with long serrated opesiae and large unfused 
condyles. Most specimens examined (dried or in 
alcohol) were of a pale blue or deep reddish colour. 

Genus Helixotionella Cook & Chimonides, 
1984b 

Zooids budded in paired interdigitating spirals. 
Avicularia with setiform mandibles. Brooding 
zooids with enlarged opesia. 

Helixotionella spiralis (Chapman, 1913) 

Selenaria marginata var. spiralis Chapman, 1913: 
184, pi. 18, fig. 33. 

Helixotionella spiralis: Cook & Chimonides, 1984b: 
257, figs 6-10, 13, 14, 17, 18, 1987: 962. 

Material Examined 

Western Australia: 32°S., 115°08'E., off Perth, 
119 m, B. Jamieson 28.viii. 1963, QM GH1 150(1); 
80 Nm (146.4 km) W of Eucla, 81 fms (148 m), 
iii.1912, SAM L561(2). 

Distribution 

Previously known from 40 km west of Jurien Bay, 
Western Australia, 137.2 m, with fossils from 
Victoria (Oligocene to Pliocene). The present 
material adds two localities to the species' range in 
southern Western Australia (one in the Great 
Australian Bight), and increases the recorded depth- 
range to 119-148 m. 

Remarks 

The two colonies in the SAM were alive when 
collected. In this species, the paired interdigitating 



THE BRYOZOAN FAMILY SELENARIIDAE 



spirals of zooids form the entire minute colony, 
which rarely exceeds 2 mm in diameter. The 
avicularian opesia is serrated and the condyles 
unfused. There is usually only one pair of basal 
avicularia. 

H. spiralis is remarkable for the extent of its 
temporal range, from Oligocene to Recent (Cook & 
Chimonides, 1987:962). 

Helixotionella scutata Cook & Chimonides, 1984b 

Helixotionella scutata Cook & Chimonides, 1984b: 
265, figs 3-5, 11, 15, 16, 19. 

Material Examined 

Western Australia: 80 Nm (146.4 km) W of 
Eucla, 81 fms (148 m), iii.1912, SAM L562(9). 

Distribution 

Previously known only by the type series from 40 
km west of Jurien Bay, south-western Western 
Australia, 137.2 m. The present material adds the 
Western Australian sector of the Great Australian 
Bight to the known distribution of this species, and 
increases its recorded depth-range to 148 m. 

Remarks 

The nine colonies examined were all alive when 
collected. H. scutata is distinguished by its 
bifurcated spiral series of zooids, and by its 
avicularia having fused condyles and an opesia 
obscured by a flattened scuta. 



Genus Selenaria Busk, 1854 

Zooids budded radially in successive zones. Central 
zone of closed zooids, surrounded by autozooids and 
avicularia. Next zone composed of enlarged, female 
brooding zooids, next zone of peripheral, non- 
feeding male zooids. Avicularia large, scattered, 
with S-shaped or reflexed condyle system, 
mandibles very long, formed from alternating discs 
of calcified and cuticular tissue. 

Selenaria bimorphocella Maplestone, 1904 

Selenaria bimorphocella Maplestone, 1904: 213, pi. 
24, fig. 3; Cook & Chimonides, 1985a: 301, figs 5d, 
15, 16. 

Selenaria punctata: Livingstone (non Tenison- 
Woods) 1928: 114. 



South Australia: 35 Nm (64 km) SW of North 
Neptunes, 104 fms (190 m), i.1905, SAM L565(3) 
E of North Neptunes, 45 fms (82 m), SAM L549(l) 
S of Troubridge L, 20 fms (36.6 m), SAM L567(2) 
Investigator Strait, 20 fms (36.6 m), SAM L416(3) 
Investigator Strait, 2 fms (3.7 m),17 fms (31 m) and 
20 fms (36.6 m) QM GH 1668(21), and no depth, 
SAM L568(l); off Point Marsden, Kangaroo I., 15 
fms (27.5 m), QM GH 1646(4); off Ardrossan, 6-8 
fms (11-14.6 m), SAM L373(l); Gulf St Vincent, 
12 fms (22 m), QM GH1673(1), 17 fms (31 m), 
SAM L566(77); Yankalilla Bay, 20 fms (36.6 m), 
SAM L569(6); Backstairs Passage, 17 fms (31 m), 
QM GH 1636(9) and no depth, SAM L570(56); 
between Backstairs Passage and The Pages, 25 fms 
(46 m), SAM L572(l); Cape Jaffa, 90 fms (165 m), 
QM GH 1639(1), 1643(1) Beachport, SAM L572(l); 
no data, SAM L573-577(32). 

Distribution 

Previously recorded from South Australia (Gulf 
St Vincent, Investigator Strait and south of Eyre 
Peninsula) and Bass Strait, 31-183 m, and as fossils 
from the Pliocene of Victoria. The present material 
extends the known range westward to the Western 
Australian sector of the Great Australian Bight, and 
adds several localities to the South Australian 
distribution, notably Cape Jaffa and Beachport in 
the South-East; it also increases the depth range to 
3.7-220 m. 



Remarks 

Of the 172 SAM colonies examined, 45 were 
alive when collected; L373 and L416 are the 
samples reported by Livingstone (1928) as S. 
punctata. A further sample identified by Livingstone 
(MS) as 5. punctata, from Yankalilla, consists of 
five colonies of 5. bimorphocella (L569) and one of 
S. concinna Teni son- Woods, 1880 (L583). 

The colonies of 5. bimorphocella are large and 
flat, with very large ancestrulae. The species is 
distinguished by the considerable sexual 
dimorphism of the zooids (the male zooids having a 
trifoliate opesia and no opesiules), and the 
autozooidal cryptocyst having a sinuate opesia. 

Pace Cook & Chimonides, 1985a: 303, S. 
bimorphocella does not replace S. punctata along 
the southern coasts of Australia, for the latter does 
occur there in small numbers (see below). However, 
to judge from the large number of samples in the 
present material, S. bimorphocella seems to be by 
far the commonest selenariid in the region. 



Material Examined 

Western Australia: 60 Nm (109.8 km) W of 
Eucla, SAM L563(l); W of Eucla, 50-120 fms (92- 
220 m), SAM L564 (1); W of Eucla, 72 fms (132 
m), QMGH1714(1). 



Selenaria punctata Tenison-Woods, 1880 

Selenaria punctata Tenison-Woods, 1880: 9, pi. 2, 
figs 8a-c; Cook & Chimonides, 1985a: 303, figs 5e, 
9, 10, 17, 19. 



S. A. PARKER & P. L. COOK 



Selenaria fenestrate. Haswell, 1880: 42. 

Selenaria partipunctata Maplestone, 1904: 214, pi. 

24, fig. 4. 

Material Examined 

Western Australia: 32°S., 115°08'E., off Perth, 
119 m, B. Jamieson 28.viii. 1963, QM GH 1150(1); 
80 Nm (146.4 km) W of Eucla, 81 fms (148 m), 
SAM L578(2); 19°32'S., 118°08'E., NW of Port 
Hedland, 50-52 m, 26.iii. 1982, WAM 4-:>2(i); 
20°19 , S., 116°47'E., off Legendre Id, 42 m, WAM 
1864-88(19); Mermaid Sound, Dampier 
Archipelago, 10.ii.1981, WAM 2-92(1); between 
Dampier and Port Hedland, WAM 87-89(1). 

South Australia: Off Cape Jaffa, 130 fms (238 
m), SAML579(1). 

Distribution 

Previously recorded from Western Australia, 
eastern Queensland and New South Wales, 11-137 
m, also as a fossil from south-western Western 
Australia (Pliocene). The present material adds 
South Australia (South-East) and the Western 
Australian sector of the Great Australian Bight to 
the known range of the species, and extends the 
lower depth-range to 238 m. 

Remarks 

As noted above, Livingstone's (1928) report of S. 
punctata from South Australia is referred to S. 
bimorphocella. The colonies of S. punctata are 
small, domed, with small ancestrulae; the 
autozooidal opesiae are D-shaped with closely- 
apposed opesiules; brooding zooids have a bar 
across the orifice, and male zooids have long, paired 
opesiules. Some of the colonies from Western 
Australia (SAM) have a diameter of only 3 mm and 
still retain frontal membranes, but few or no 
mandibles, many of the others (WAM) are worn. 
The colony from South Australia is very worn. 

The widespread S. punctata is very similar to 5. 
parapunctata Cook & Chimonides, 1985a, which 
appears to replace it in Bass Strait (see Discussion). 
5. parapunctata differs by its widely spaced 
opesiules, absence of an orificial bar in brooding 
zooids, and S-shaped, rather than reflexed 
avicularian condyles. 

Selenaria pulchella MacGillivray, 1895 

Selenaria squamosa var. pulchella MacGillivray, 
1895:48, pi. 7, fig. 13. 

Selenaria pulchella: Cook & Chimonides, 1984b: 
262; 1985a: 307, figs 4, 20. 

Material Examined 

Western Australia: 80 Nm (146.4 km) W of 
Eucla, 81 fms (148 m), SAM L580(3). 



Distribution 

Previously recorded from Jurien Bay, 137 m, 
south-western Western Australia, with fossils from 
the Miocene of Victoria. The present sample adds 
the Western Australian sector of the Great 
Australian Bight to the species' known range, and 
increases the depth-range to 148 m. 

Remarks 

Of the three colonies examined, two were alive 
when collected. The colonies of this species are 
minute (2-3 mm in diameter). The zooids have 
small lateroproximal opesiular indentations, the 
male zooids possess small paired opesiules, and the 
avicularia have punctate frontal shields. All three 
present colonies were sexually mature, with male 
zooids and large peripheral avicularia, though no 
mandibles were present. 

Selenaria concinna Tenison-Woods, 1880 

Selenaria concinna Tenison-Woods, 1880: 10, pi. 2, 
figs lla-c; Cook & Chimonides, 1987: 950, figs 1, 
11,24,28, 30. 

Material Examined 

Western Australia: 80 Nm (146.4 km) W of 
Eucla, 81 fms (148 m), SAM L581(l). 

South Australia: 35 Nm (64 km) SW of North 
Neptunes, 104 fms (190 m), i.1905, SAM L582(2); 
Yankalilla Bay, 20 fms (36.6 m), SAM L583(l); 
Backstairs Passage, SAM L584(5); Cape Jaffa, 90 
fms (165 m), QMGH1643(3). 

Distribution 

Previously recorded from eastern Queensland, 
New South Wales, Bass Strait and New Zealand, 
33-148 m, and as a fossil from the Pliocene of 
Western Australia and the Miocene of Victoria. The 
present material adds South Australia and Western 
Australia, to the species' known range, and extends 
the lower depth-limit to 190 m. 

Remarks 

One of a complex of five very similar species 
(Cook & Chimonides, 1987), S. concinna is 
distinguished by its avicularia, which have a 
serrated proximal opesia and a distal calcified 
bridge. None of the present specimens is larger than 
7 mm in diameter, and all are slightly worn. 

Selenaria varians Cook & Chimonides, 1987 

Selenaria varians Cook & Chimonides, 1987: 957, 
figs 2, 32, 33. 

Material Examined 

South Australia: 35 Nm (64 km) SW of North 



THE BRYOZOAN FAMILY SELENARIIDAE 



Neptunes, 104 fms (190 m), i.1905, SAM L585(3); 
Cape Willoughby, Kangaroo I., 23 fms (42 m), QM 

GH3160(1). 

Distribution 

Previously recorded from Bass Strait and New 
South Wales, 46-95m. The present material adds 
South Australia to the known distribution of the 
species, and amplifies the depth-range to 42-190 m. 

Remarks 

The four colonies examined have a maximum 
diameter of 4 mm and are all slightly worn. 5. 
varians is distinguished by its autozooidal opesiae 
becoming proportionately larger with astogeny. The 
avicularia have a marginally serrate opesia. 

Selenaria exasperans Cook & Chimonides, 1987 

Selenaria exasperans Cook & Chimonides, 1987: 

957, figs 5, 12,34,35. 

Material Examined 

South Australia: 35 Nm (64 km) SW of the 

North Neptunes, 104 fms. (190 m), i.1905, SAM 
L586(2); Cape Jaffa, 130 fms (238 m), SAM 
L587(l); Cape Jaffa, 90 fms (165 m), QM 
GH1639(2); Beachport, 110 fms (201 m), SAM 

L588(2). 



Distribution 

Previously recorded from Bass Strait and New 
South Wales, 79-148 m. The present series adds 
South Australia to the known distribution, and 
extends the depth-range to 238 m. 

Remarks 

S. exasperans is distinguished by the presence of 
a proximo- and disto-lateral avicularium beside the 
ancestrula, a pattern unique in the genus. 
Autozooidal opesiae are D-shaped, and avicularian 
opesiae serrate. All five colonies examined 
(maximum diameter 4 mm) are slightly worn, but 
clearly show the ancestrula and adjacent avicularia 
and autozooids. 



Selenaria hexagonalis Maplestone, 1904 
(Fig. 1,A) 

Selenaria hexagonalis Maplestone, 1904 (part): 214, 
pi. 24, fig. 5; Cook & Chimonides, 1987 (part): 948, 
figs 10, 20, 21 (not figs 8, 22, 23, = S. verconis sp. 
nov.) 

Material Examined 

Western Australia: King George Sound, 28 fms 
(51 m), SAM L589(l); no data, SAM L590- 

592(22). 





FIGURE 1. Sketches of zooids of Selenaria; M, male, F, female, Z, autozooid, A, avicularium. A, S. hexagonalis Maplestone. 
B, S. verconis sp. nov. Scale = 0.50 mm. 



S. A. PARKER & P. L. COOK 



South Australia: Investigator Strait, 15 fms (27,5 
m), NMV P62753, lectotype (selected by Cook & 
Chimonides, 1987: 948). 

Distribution 

Known from King George Sound, south-western 
Western Australia, 51 m and Investigator Strait, 
South Australia, 27.5 m. 

Remarks 

Colonies large (diameter 13-15 mm when 
sexually mature), robust and thickened basally. 
Autozooids hexagonal in frontal view, with 
subcentral, almost circular opesia. Centre of 
operculum just distal to centre of opesia. 
Subperipheral brooding zooids large, with elongated 
opesia, raised distally. Peripheral male zooids with 
very large, oval opesia, without subopercular 
tubercle. Avicularian shield with 12-16 stout bars, 
fused medially; condyle system not much reflexed, 
S-shaped. 

The distinctness of this species from the next (S. 
verconis sp. nov.) was suspected by Cook & 
Chimonides (1987). 5. hexagonalis sensu stricto has 
a more restricted range than S. verconis sp. nov., 
being known with certainty from only one district in 
Western Australia and one in South Australia. 

One very large colony (one of the nine in SAM 
L591), 14 mm in diameter, had apparently ceased 
growing when 1 1 mm in diameter, just before sexual 
maturity. Subsequent growth around the perimeter 
is marked by swollen basal calcification and 
concentric series of brooding and male zooids, 
interspersed with large avicularia. The avicularia are 
orientated radially, and their condyle systems seem 
normal, but the frontal shield of bars of calcification 
is orientated at right angles to the normal direction. 
The setiform mandibles would not have been 
affected by the orientation of the frontal shield. The 
two other mature colonies in this sample show no 
abnormalities. One of the three colonies in sample 
SAM L592 has large acrothoracid cirripede burrows 
in the central area. 



Selenaria verconis sp. nov. 
(Fig. 1,B) 

Selenaria hexagonalis Maplestone, 1904 (part): 214 
(Jimmy's Point Farm, Victoria, Pliocene); Cook & 
Chimonides, 1987 (part): 948, figs 8, 22, 23. 

Material Examined 

HOLOTYPE: Queensland: Off Townsville, 2.5- 
12 m, coll. Cook & Chimonides, 1982, BMNH 
1984. 12.24.14 (illustrated by scanning electron 
micrographs in Cook & Chimonides, 1987, figs 8, 
22, 23). 

PARATYPES: Western Australia: King George 



Sound, 22-28 fms (40-51 m), SAM L593(l), SAM 
L594(l); ditto, 28 fms (51 m), SAM L594 (1); 40 
Nm (73.2 km) W of Eucla, 75-105 fms (137-192 
m), iii.1912, SAML596(1). 

South Australia: Off Adelaide, 36-64 m, BMNH 
1928.9.13.78. 

Queensland: Port Denison, BMNH 
1984.12.24.36. 

Victoria (Pliocene): Jimmy's Point Farm, coll. 
Maplestone. NMV T1757. (Last three samples and 
holotype listed under S. hexagonalis by Cook & 
Chimonides, 1987:948). 

Description 

Colonies 10-11 mm when sexually mature. 
Autozooids with oval or suboval opesia, proximal 
edge sometimes almost straight; cryptocyst finely 
serrated. Operculum dark, placed above the distal 
half of the opesia. Subperipheral brooding zooids 
large, raised distally. Peripheral male zooids with 
elongated oval or pyriform opesia; subopercular 
tubercle absent. Avicularian shield with 14-20 fine 
bars, fused medially; frontal area narrowing distally, 
flanked by cryptocystal margins of adjacent zooids; 
condyle system reflexed. 

Etymology 

Verconis, L., genitive singular of Verco 
(construing the surname as a noun of the Third 
Declension); named after Dr Sir Joseph Cooke 

Verco (1851-1935). 

Distribution 

Known from south-western Western Australia 
(King George Sound), South Australia (Gulf St 
Vincent) and eastern Queensland, 2.5-192 m, and 
as a fossil from the Pliocene of Victoria. 

Remarks 

When redescribing S. hexagonalis. Cook & 
Chimonides (1987) noted the wide range of 
character states shown by both Maplestone's 
specimens and those in the BMNH, concluding that 
two partly sympatric forms might be involved. The 
four colonies from Western Australia examined 
here, some of which were collected with S. 
hexagonalis s. s. y have the unequivocal character 
correlations of the Queensland specimens. This 
indicates, as previously suspected, that two distinct 
and sympatric species are present, one (5. 
hexagonalis) with a restricted distribution the other 
(S. verconis) with a wider geographical range and a 
fossil record. 

5. verconis most resembles S. hexagonalis 
Maplestone, differing by its less robust zoaria, 
smaller and proportionately longer zooids with oval 
to suboval opesiae and more distal opercula, and by 
the avicularia having 14-20 fine bars (vs 12-16 



THE BRYOZOAN FAMILY SELENARIIDAE 



stout bars in S. hexagonalis) and a more rellexed 
condyle system (see Fig. IB). 

'Selenaria' alata auctt. 

Selenaria alata: Cook & Chimonides, 1985c: 339, 
figs 1, 2, 5, 9, 11-13 (Recent material only). Non 
Selenaria alata Tenison-Woods, 1880: 1 1, pi. 2, figs 
12a-c. 

Material Examined 

South Australia: 35 Nm (64 km) SW of the 
North Neptunes, 104 fms (190 m), i.1905, SAM 
L597(3); no data, SAM L598(l). 

Distribution 

Known previously only from Bass Strait, 46-95 
m. The present material adds South Australia to the 
recorded range of the species, and increases the 
lower depth-limit to 190 m. 

Remarks 

Of the four colonies examined, one was alive 
when collected. This species is characterized by its 
large, asymmetrical, unfused avicularian condyles 
and large trifoliate autozooidal opesiae. 

The Recent colonies from Bass Strait were all 
regenerated from fragments, and differed slightly 
from fossil S. alata in their less trifoliate autozooid 
opesiae. The present specimens do not exceed 12 
mm in diameter, and two have mandibles 
resembling those of Otionella squamosa, see Cook 
& Chimonides (1985c, fig. 1 1). The ancestrulae are 
all worn or partly obscured, but do not appear to be 
as large as those of the fossil colonies. The 
autozooidal opesiae resemble those of the Bass 
Strait population, and are only slightly trifoliate. 
The somewhat ambiguous character of Recent S. 
alata audi., which shares features with both fossil 
S. alata and S. lata, will be discussed in a later 
article (Bock & Cook in prep.). The character of the 
avicularian condyles and mandibles of S. lata, S. 
alata and S. alata auctt., together with the absence 
of zones of distinctive brooding and male zooids, 
means that none of these three species can be 
referred to Selenaria s. s., and they require a generic 
grouping of their own. 



Discussion 

The present collections, though sparse in 
comparison with those previously reported from 
Bass Strait and New Zealand (Cook & Chimonides, 
1987 and references therein; Nelson et al. 1988), 
are nevertheless significant in that they allow a first 
estimate of the diversity of the Selenariidae 
occurring in southern Western Australia and South 
Australia. The 16 species listed constitute a fairly 



diverse fauna, comparable to that of Bass Strait with 
its 18 species. New State records are: Western 
Australia: Selenaria bimorphocella, S. concinna, S. 
hexagonalis and S. verconis; South Australia: 
Otionella nitida, O. australis, S. punctata, S. 
concinna, S. varians, S. exasperans, S. verconis and 
'S'. alata auctt. Whereas most of the new locality 
records constitute only minor range-extensions (e.g. 
S. concinna), the extension in some cases is 
considerable. For example, Helixotionella spiralis, 
H. scutata and S. pulchella, previously known in 
the Recent only from the Jurien Bay district of 
Western Australia, are reported from the Eucla 
district of the Great Australian Bight, some 1 700 
km to the south-east. Sufficient samples have now 
been analyzed from Bass Strait to permit the 
suggestion that these last three species are today 
genuinely absent from that region (though H. 
spiralis and S. pulchella have a fossil record in 
Victoria), and that the Great Australian Bight may 
well mark the eastern limit of their modern 
distribution. In addition, 5. hexagonalis is reported 
from the Albany district of Western Australia, over 
2 000 km west of its previous western limits in 
South Australia, and 5. varians and S. exasperans 
are shown to occur at St Francis I., 1 000 km west 
of their original localities in Bass Strait (with 5. 
exasperans occurring also at the intermediate 
localities of Cape Jaffa and Beachport in south- 
eastern South Australia). 

The distribution of the species-pair S. punctata 
and S. parapunctata is of particular interest. The 
former is now known from most parts of the 
Australian continental shelf except Bass Strait, the 
Northern Territory and northern Queensland. Its 
absence from relatively well-collected Bass Strait 
may be real; moreover, in Bass Strait it appears to 
be replaced by S. parapunctata, which so far has 
not been reported from elsewhere. 

Of the 18 species currently known from Bass 
Strait, seven are still notably absent from collections 
made further west: these are Otionella minuta, O. 
auricula, Selenaria initia, S. minor, S. maculata, S. 
kompseia, and 5. maplestonei (see Cook & 
Chimonides, 1985a, b; 1987). It is possible that at 
least some of these will eventually be found in 
South Australia and Western Australia. After the 
discovery of living H. spiralis, S. pulchella, S. initia 
and S. minor (originally described as fossils), it is 
possible that, with further collecting, other fossil 
species will also be discovered to be extant. 

Acknowledgments 

We thank Mary Spencer Jones (BMNH), John Hooper 
(QM) and Diana Jones (WAM) for information on specimens 
in their care, and Philip Bock (Royal Melbourne Institute of 
Technology) for examining and identifying for us the QM's 
holdings of selenariids. 



10 



S. A. PARKER & P. L. COOK 



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COOK, P. L. & CHIMONIDES, P. J. 1984b. Op. cit. 2. 
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COOK, P. L. & CHIMONIDES, P. J. 1985a. Op. cit. 3. 
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COOK, P. L. & CHIMONIDES. P. J. 1985b. Op. cit. 4. 
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COOK, P. L. & CHIMONIDES, P. J. 1985c. Op. cit. 5. 
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COOK, P. L. & CHIMONIDES, P. J. 1986. Op. cit. 6. 
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THE BRYOZOAN FAMILY SELENARIIDAE 



11 



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A CHECKLIST OF HELMINTH PARASITES OF AUSTRALIAN AMPHIBIA 

Diane P. Barton 

Summary 

This checklist includes all original references, and any other references which do more than repeat 
original work, of helminths occurring in Australian amphibians published up to 1992. Museum 
listings are also included, where available. Most records pertain to free-ranging animals; where they 
do not, they have been annotated appropriately. 



A CHECKLIST OF HELMINTH PARASITES OF AUSTRALIAN AMPHIBIA 



DIANE P. BARTON 



BARTON, D. P. 1994. A checklist of helminth parasites of Australian Amphibia. Rec. S. Aust. Mm. 
27(1): 13-30. 

This checklist includes all original references, and any other references which do more than repeat 
original work, of helminths occurring in Australian amphibians published up to 1992. Museum listings 
are also included, where available. Most records pertain to free-ranging animals; where they do not, they 
have been annotated appropriately. 

Helminths are arranged as follows: Monogenea, Digenea, Cestoda, Nematoda, Acanthocephala, in 
both the parasite-host and host-parasite checklists. 

Hosts are presented by family with consideration given to recent taxonomic changes. 

D. P. Barton, Zoology Department, James Cook University, Townsville, Queensland, Australia, 4811. 
Manuscript received 20 August 1992. 



Introduction 

In 1939 May Young produced a checklist of 
helminth parasites recorded from Australian hosts. 
Thirteen amphibian hosts, infected with a total of 
30 helminth species, were included. There has been 
no further compilation of solely Australian records 
from amphibians since then. The aim of this work is 
to produce an updated checklist of amphibian 
helminth parasites in Australia. 

Included in this checklist are all original 
references, and any other references which do more 
than repeat original work, published up to 1992. 
Museum collections of amphibian parasites are also 
included, where available. Most records pertain to 
free ranging animals; where they do not, they have 
been annotated appropriately (e.g. experimental). 



Comments on host taxonomy 

Host names used in this checklist follow Cogger 
(1992), with the following exceptions: 

i) Kyarranus Moore, 1958 is accepted as a valid 
genus (see Frost 1985). 

ii) Litoria pearsoniana (Copland, 1961) is 
accepted as a valid species (see Frost 1985). 

In the parasite-host checklist, host names are 
given as they were listed in the original publications. 
In the host-parasite checklist, the names have been 
updated to those used by Cogger (1992). The 
original names are also given with reference to the 
new name when there is an element of confusion. 

All species formerly referred to the family 
Leptodactylidae are now placed in the family 
Myobatrachidae (see Cogger 1992). 

All Hyla species are now referred to the genus 
Litoria (see Cogger 1992). 

Limnodynastes dorsalis and L. d. dumerilii 



recorded from South Australia, Queensland and 
New South Wales are referred to L. dumerilii. 
Limnodynastes dorsalis is only present in Western 
Australia (see Cogger 1992). 

Litoria aurea and L. a. raniformis recorded from 
South Australia, Victoria and Tasmania are referred 
to L. raniformis (see Cogger 1992). 

Litoria aurea recorded from Western Australia is 
referred to Litoria spp., as the range of L. aurea 
does not extend to Western Australia (see Cogger 
1992). 

Crinia sp. recorded from the Flinders Ranges, 
South Australia, in the Australian Helminthological 
Collection (AHC) list are most likely C. riparia. A 
group of these frogs was collected from Warren 
Gorge, which is within the range of C. riparia (Dr 
Margaret Davies, pers. comm.). A more precise 
geographical location is, however, required to 
differentiate C. riparia from C. signifera, so they 
must remain Crinia sp. 

Litoria jervisiensis recorded from South Australia 
is referred to L. ewingii (see Cogger 1992). 

Mixophyes sp. collected from the Bunya Mts, 
Queensland (AHC 6172), could be either M. 
fasciolatus or M. iteratus. Examination of the host 
specimen would be needed to determine the exact 
species. 

Uperoleia marmorata collected from New 
England National Park, New South Wales (AHC 
8055), could be either U. rugosa or U. laevigata. 
Davies & Littlejohn (1986) showed both species to 
be present in this region, while U. marmorata was 
restricted to the north-west of Western Australia. 
All U. marmorata in this checklist are from eastern 
Australia and helminths from these host specimens 
are referred to Uperoleia spp. Again, examination 
of the host specimen would be needed to determine 
which species is correct. 

Bufo marinus was introduced to Australia in 1935 



14 



DIANE P. BARTON 



from South America (via Hawaii) (Easteal 1981). It 
is 'naturally' found in Queensland, northern New 
South Wales and eastern Northern Territory. Any 
locations recorded out of this range are from 
laboratory animals acquired from a commercial 
supplier. 

Records of helminths from frogs in New Guinea 
are included only if that frog species is also found in 
Australia. 



Comments on helminth taxonomy 

Helminth nomenclature follows Prudhoe & Bray 
(1982) for Monogenea, Digenea and Cestoda, the 
CIH Keys to the Nematode Parasites of Vertebrates 
(Hartwich 1974; Chabaud 1975a, 19756, 1978; 
Anderson & Bain 1976, 1982; Petter & Quentin 
1976; Durette-Desset 1983) and Spencer Jones & 
Gibson (1987) for the Nematoda, and Amin (1985) 
for the Acanthocephala. 

The taxonomic status of many genera and species 
infecting amphibians is in need of revision. 
Generally, original records of helminths are treated 
as correct, unless it is known to the author that 
appropriate revision has taken place. 

All helminths are recorded in the parasite-host 
checklist under the current name with any synonyms 
also listed. 

All previous records of lung nematodes from 
Australian frogs have been referred to Rhabdias 
hylae Johnston & Simpson, 1942, by Ballantyne 
(1971), a view accepted in this checklist. 

Nasir & Diaz (1971) synonymised the Australian 
representatives of the genus Mesocoelium as M. 
megaloon Johnston, 1912, and M. monas (Rudolphi, 
1819), Teixeira de Freitas, 1958 (M. microon 
Nicoll, 1914, M, mesembrinum Johnston, 1912, M. 
oligoon Johnston, 1912). This is not accepted here, 
pending further work on the genus. 

Frogs often serve as an intermediate host for 
cestodes, being infected with plerocercoids/spargana 
in the musculature. The identification of this life 
cycle stage is impossible without knowledge of the 
definitive host. In other groups (Acanthocephala, 
Digenea), larval stages are often identifiable. 



Explanation of Format 

This checklist has been compiled from all 
published records up to 1992 known to the author, 
and from lists of museum holdings. 

References to Prudhoe & Bray (1982) on 
microfiche are shown as 'mf following the page 
number. 

The lists- are arranged as follows: 
1. Parasite species are arranged systematically. The 



amphibian hosts are listed for each helminth 
followed by the state or territory of origin (? 
denotes the state or territory was not referred to), 
literature references, and museum collection 
numbers, where available. The hosts are 
arranged with the type host first and all others 
listed alphabetically after. 

2. Host species are arranged alphabetically within 
each family. The helminths from each host 
species are listed below the host with the phase 
of development and site of infection recorded, 
where known. 

3. References. 

Authors whose names appear frequently are 
referred to, where appropriate, by initials, as 
follows: 

LMA L. Madeline Angel 
MRY May R. Young 
PMM Patricia M. Mawson 
SJJ Stephen J. Johnston 
THJ T. Harvey Johnston 

The major helminth parasite groups are referred 
to by their initials: 
M Monogenea 
D Digenea 
C Cestoda 
N Nematoda 

A Acanthocephala 

Museums and other sources which are referred to 
as having amphibian parasites in their collection are 
abbreviated as follows: 
AHC Australian Helminthological Collection, 

(in the South Australian Museum) 

Adelaide, SA 
AM Australian Museum, Sydney, NSW 

BM(NH) Natural History Museum, London, 

England 
CAS Institute of Parasitology, Czechoslovak 

Academy of Sciences, Ceske Budejovice, 

Czechoslovakia 
QM Queensland Museum, Brisbane, Qld 

SAM South Australian Museum, Adelaide, SA 
SP Personal collection of Ms Sylvie Pichelin, 

Parasitology Department, University of 

Queensland, Brisbane, Qld 
TM Tasmanian Museum and Art Gallery, 

Hobart, Tas 

State names are abbreviated as follows: 
NSW New South Wales 
NT Northern Territory 

Qld Queensland 

SA South Australia, including Kangaroo 1. & 

Pearson I. 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



15 



Tas Tasmania, including Bass Strait Islands 

(King & Flinders) 

Vic Victoria 

WA Western Australia 



Order and arrangement of parasites as presented 
under each host 

1. Phylum Platyhelminthes 

Class Monogenea Cams. 1863 

Order Polyopisthocotylea Odhner, 1912 

Family POLYSTOMATIDAE Carus, 1863, 

emended Gamble, 1 896 

Subfamily Polystomatinae Gamble, 1896 

Class Trematoda Rudolphi, 1808 
Order Digenea Van Beneden, 1858 
Suborder Prosostomata Odhner, 1905 

Family PARAMPHISTOMATIDAE Fischoeder, 

1901 

Subfamily Diplodiscinae Cohn, 1904 

Family GORGODERIDAE Looss, 1901 

Family ALLOCREADIIDAE Stossich, 1903 

Family PLAGIORCHIIDAE Liihe, 1901, 
emended Ward, 1917 

Subfamily Haematoloechinae Teixeira de Freitas 
& Lent, 1939, emended Yamaguti, 1958 

Family TELORCHIIDAE Stunkard, 1924 
Subfamily Opisthioglyphinae Dollfus, 1949 

Family BRACHYCOELIIDAE Johnston, 1912 

Family LECITHODENDRIIDAE Odhner, 1910 

Family BRACHYLAIMIDAE Joyeux & Foley, 
1930 

Family DIPLOSTOMIDAE Poirier, 1886 

Family DOLICHOPERIDAE Yamaguti, 1971 

Not further identified 

Class Cestoidea Rudolphi, 1808 
Order Pseudophyllidea Carus, 1 863 

Family DIPHYLLOBOTHRIIDAE Liihe, 1910 

Order Proteocephalidea Mola, 1928 

Family PROTEOCEPHALIDAE La Rue, 1911 

Order Cyclophyllidea Braun, 1900 

Family NEMATOTAENIIDAE Ltihe, 1910 

Not further identified 



2. Phylum Nematoda 

Class Secernentea 
Order Rhabditida 

Superfamily Rhabditoidea 

Family RHABDIASIDAE Railliet, 1916 

Order Strongylida 

Superfamily Trichostrongyloidea 

Family MOLINEIDAE (Skrjabin & Schulz, 1937) 

Durette-Desset & Chabaud, 1977 

Order Oxyurida 

Superfamily Oxyuroidea 

Family PHARYNGODONIDAE Travassos, 1919 

Order Ascaridida 

Superfamily Cosmocercoidea 

Family COSMOCERCIDAE (Railliet, 1916, 

subfam.) Travassos, 1925 

Superfamily Ascaridoidea 

Family ASCARIDIDAE Baird, 1853 

Order Spirurida 

Superfamily Physalopteroidea 

Family PHYSALOPTERIDAE (Railliet, 1893, 

subfam.) Leiper, 1908 

Superfamily Habronematoidea 
Family HEDRURIDAE Railliet, 1916 

Superfamily Filarioidea 
Not further identified 



3. Phylum Acanthocephala 

Class Palaeacanthocephala Meyer, 1931 

Order Echinorhynchida Southwell & MacFie, 1925 

Family ECHINORHYNCHIDAE Cobbold, 1876 

Order Polymorphida Petrochenko, 1956 

Family PLAGIORHYNCHIDAE Golvan, 1960 

Not further identified 



Order and arrangement of hosts as presented in 
host-parasite checklist 

Class Amphibia 
Order Anura 

Family MYOBATRACHIDAE 

Adelotus 

Arerophryne 



16 



DIANE P. BARTON 



Assa 

Crinia 

Geocrinia 

Heleioporus 

Hyperoleia 

Kyarranus 

Leptodactylid 

Limnodynastes 

Metacrinia 

Mixophyes 

Myobatrachid 

Neobatrachus 

Paracrinia 

Philoria 

Pseudophryne 

Ranidella 

Rheobatrachus 

Taudactylus 

Uperoleia 

Family HYLIDAE 

Chiroleptes 

Cyclorana 

Hyla 

Litoria 

Family RANIDAE 
Rana 

Family BUFONIDAE 
Bufo 

Unidentified Anura 



Parasite-host checklist 
1. Phylum Platyhelminthes 

Class Monogenea Cams. 1863 

Order Polyopisthocotylea Odhner, 1912 

Family POLYSTOMATIDAE Cams, 1863, 

emended Gamble, 1 896 

Subfamily Polystomatinae Gamble, 1 896 

Parapoly stoma bulliense (Johnston, 1912), 

Ozaki, 1935 

syn. Polystomum bulliense Johnston, 1912 
Hyla phyllochroa, NSW, SJJ 1912: 297, AM 
W.346, QM GL 12109, GL 12160, AHC 2200 
(wholemount), 2217-2219 (sections) 
Hyla lesueurii, NSW, SJJ 1912: 297 
Litoria citropa, NSW, AHC 5167 
Litoria pearsoniana, Qld, SP 

Parapoly stoma sp. 

Litoria nyakalensis, Qld, SP 

Class Trematoda Rudolphi, 1 808 
Order Digenea Van Beneden, 1858 
Suborder Prosostomata Odhner, 1905 



Family PARAMPHISTOMATIDAE Fischoeder, 

1901 

Subfamily Diplodiscinae Cohn, 1904 

Diplodiscus megalochrus Johnston, 1912 

Hyla aurea, NSW, SJJ 1912: 302, AM W.332, 

QMGL 11851 

Frog, NSW, AHC 3310 

Hyla caerulea, Qld, THJ 191 66: 60 

Limnodynastes peronii, NSW, SJJ 1912: 302 

Litoria caerulea, Qld, Prudhoe & Bray 1982: 

199 mf 

Diplodiscus microchrus Johnston, 1912 

Hyla ewingii, NSW, SJJ 1912: 307, AM W.333 
Limnodynastes tasmaniensis, NSW, SJJ 1912: 
307 

Diplodiscus sp. 

Bufo marinus, Qld, AHC 14, 2978, 3028, 

3553, 3563, 3576, 3875 

Hyla aurea, NSW, AHC 12683 

Hyla caerulea, Qld, QM GL 12350 

Amphistome 

Bufo marinus, Qld, AHC 4944 

Distoma sp. 

Hyla aurea, ?, MRY 1939: 74 

Family GORGODERIDAE Looss, 1901 

Gorgodera australiensis Johnston, 1912 
Hyla aurea, NSW, SJJ 1912: 326, AM 
W.340a, AM W. 395, AM W. 19850, QM GL 
11860, GL 12161 

Limnodynastes dorsalis, SA, AHC 351 1 
Limnodynastes peronii, NSW, SJJ 1912: 326, 
AM W.340 (this number is given for H. aurea 
in SJJ 1912: 326, but in AM records is for L. 
peronii) 

Gorgodera sp. 

Hyla aurea, NSW, AHC 12680; Vic, AHC 

4539; SA, AHC 3529, 3532 

Limnodynastes dorsalis, SA, AHC 3498, 3502, 

12698 

Limnodynastes tasmaniensis, SA, AHC 3489 

Family ALLOCREADIIDAE Stossich, 1903 

Allocreadiidae sp. 

Cyclorana cultripes, Qld, QM GL 1 1285 

Family PLAGIORCH1IDAE Liihe, 1901, 
emended Ward, 1917 

Subfamily Haematoleochinae Teixeira de Freitas 
& Lent, 1939, emended Yamaguti, 1958 

Haematoleochus australis (S.J. Johnston, 1912), 
Inglis, 1932 syn. Pneumonoeces australis S.J. 
Johnston, 1912 

Hyla aurea, NSW, SJJ 1912: 321, AM W.339, 
W.339a, W.396, W. 19849; ?, QM GL 11868, 
GL1197 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



17 



Limnodynastes peronii, NSW, SJJ 1912: 321 
Litoria aurea, Tas, AHC 5404 
Litoria moorei, WA, Prudhoe & Bray 1982: 83 
mf, BM(NH) 1967.10.23.7-9 

Family TELORCHIIDAE Stunkard, 1924 
Subfamily Opisthioglyphinae Dollfus, 1949 

Dolichosaccus anartius (S.J. Johnston, 1912) 

Yamaguti, 1958 

syn. Brachysaccus anartius S.J. Johnston, 1912 

Hyla aurea, NSW, SJJ 1912: 317, AM W.337, 

W.398, QM GL 1 1846, AHC 12685, 12686; ?, 

QMGL 11868, GL 11997 

Limnodynastes peronii, NSW, SJJ 1912:317 
Dolichosaccus diamesus S.J. Johnston, 1912 

Hyla freycineti, NSW, SJJ 1912: 315, AM 

W.336, W.19848 
Dolichosaccus ischyrus S.J. Johnston, 1912 

Limnodynastes dorsalis, NSW, SJJ 1912: 314, 

AM W.335 

Hyla caerulea, NSW, SJJ 1912: 314; Qld, THJ 

1916:60 
Dolichosaccus juvenilis (Nicoll, 1918), 
Travassos, 1930 
syn. Brachysaccus juvenilis Nicoll, 1918 

Chiroleptes brevipalmatus, Qld, Nicoll 1918: 

368 

Cyclorana cultripes, Qld, QM GL 1 1 280 
Dolichosaccus symmetrus (SJ. Johnston, 1912), 
Yamaguti, 1958 
syn. Brachysaccus symmetrus Johnston, 1912 

Hyla caerulea, NSW, SJJ 1912: 319, AM 

W.338 

Bufo marinus, Qld, AHC 1 3 
Dolichosaccus trypherus S.J. Johnston, 1912 

Limnodynastes peronii, NSW, SJJ 1912: 310, 

AM W.334, QM 

GL 11850 

Hyla aurea, NSW, SJJ 1912: 310, QM GL 

1 1850; SA, AHC 12704 

Limnodynastes dorsalis, SA, AHC 12699 

Limnodynastes tasmaniensis, SA, AHC 3485, 

3487, 3488 

Litoria moorei, WA, BM(NH) 1968.4.19.16 
Dolichosaccus sp. 
syn. Brachysaccus sp. 

Bufo marinus, Qld, AHC 18, 2973, 2975, 

3559, 3874, 4952, 4953, 5192 

Hyla aurea, NSW, AHC 3527, 12682 

Hyla caerulea, Qld, AHC 12690 

Hyla sp., ?, MRY 1939:75 

Limnodynastes dorsalis, SA, AHC 3512, 3513, 

12677 

Limnodynastes fletcheri, SA, AHC 4583 

Limnodynastes tasmaniensis, SA, AHC 3485, 

3487, 3488 



Family BRACHYCOELIIDAE Johnston, 1912 

Mesocoelium megaloon S.J. Johnston, 1912 
Hyla ewingii, NSW, SJJ 1912: 335, AM W.343 
Litoria caerulea, ?, Freitas 1963: 179 (noted 
that this specimen should be M. mesembrinum) 
Litoria ewingii, ?, Prudhoe & Bray 1982:117 
mf 

Mesocoelium mesembrinum S.J. Johnston, 1912 
Hyla caerulea, NSW, SJJ 1912: 330, AM 
W.341 , W.341b, W.393, W.394, AHC 4538 
Bufo marinus, Qld, Yuen 1965: 271 
Litoria aurea, ?, Prudhoe & Bray 1982: 1 17 
Litoria caerulea, Qld, THJ 191 6b: 60; NSW, 
QMGL 11861 

Mesocoelium microon Nicoll, 1914 

Litoria caerulea, Qld, Nicoll 1914: 339, QM 
GL 11131 

Cyclorana cultripes, Qld, QM GL 1 1278 
Litoria gracilenta, Qld, Nicoll 1914: 339, QM 
GL 11169 

Mesocoelium oligoon S.J. Johnston, 1912 
Hyla citropus, NSW, SJJ 1912: 336 AM 
W.342 

Mesocoelium sp. 

Bufo marinus, Qld, Freeland et al. 1986: 496, 

AHC 16, 17, 2967, 2973, 2975, 3138, 3876, 

4949, 4951, 4955; SA, AHC 4547 

(Mesocoelium sp. 2 of LMA) 

Hyla caerulea, Qld, AHC 3517-3521, 3523, 

3524 

Family LECITHODENDRIIDAE Odhner, 1910 

Pleurogenoides freycineti (S.J. Johnston, 1912), 

Travassos, 1930 

syn. Pleurogenes freycineti Johnston, 1912 

Hyla freycineti, NSW, SJJ 1912: 342, AM 

W.344 
Pleurogenoides solus (S.J. Johnston, 1912), 
Travassos, 1930 
syn. Pleurogenes solus Johnston, 1912 

Hyla aurea, NSW, SJJ 1912: 345, AM W.345, 

W.19851, W. 19852 
Pleurogenes spp. 

Hyla spp., ?, MRY 1939:75 
Lecithodendriid sp. 

Bufo marinus, Qld, Freeland et al. 1986: 496 

Family BRACHYLAIMIDAE Joyeux & Foley, 
1930 

Zeylanurotrema spearei Cribb & Barton, 1991 
Bufo marinus, Qld, Cribb & Barton 1991: 207, 
QM GL 1273, 1274-76, AHC 18984, BM(NH) 
1990.12.7.3-5 

Family DIPLOSTOMIDAE Poirier, 1886 
Fibricola intermedius (Pearson, 1959), 



18 



DIANE P. BARTON 



Sudarikov, 1961 

syn. Neodiplostomum intermedium Pearson, 1959 

Hyla pearsoni, ?, diplostomula, Pearson 1961 : 

135 

Hyla caerulea, paratenic host, ?, Pearson 1961: 

136 

Hyla latopalmata tadpole, ?, Pearson 1961: 

135 

Leptodactylid sp., ?, Pearson 1961: 135 

Mixophyes fasciolatus tadpole, ?, Pearson 

1961: 135 

Family DOLICHOPERIDAE Yamaguti, 1971 

Dolichoperoides macalpini (Nicoll, 1918), 

Johnston & Angel, 1940 

syn. Dolichopera macalpini Nicoll, 1918 

Limnodynastes sp. tadpole, SA, metacercaria, 
THJ & Angel 1940: 381, AHC 201320 
Hyla aurea raniformis, SA, metacercaria, THJ 
& Angel 1940:382 

Limnodynastes dorsalis (dumerili), SA, 
metacercaria, THJ & Angel 1940: 382 
Limnodynastes tasmaniensis (platycephalus), 
SA, metacercaria, THJ & Angel 1940: 382 
Tadpole, SA, metacercaria, AHC 2725 

Digenea Not Further Identified 

Cercaria ameriannae T.H. Johnston & 

Beckwith, 1947 

Limnodynastes sp., SA, diplostomula, 
(experimental), THJ & Beckwith 1947: 578, 
AHC 20219 

Tadpole, SA, diplostomula, (experimental), 
AHC 2272 

Cercaria angelae T.H. Johnston & Simpson, 

1944 
Limnodynastes tasmaniensis tadpole, SA, 
cysts, AHC 2825; experimental infection of L. 
tasmaniensis tadpoles produced Tetracotyle 
cysts (THJ & Simpson 1944: 131) 
Tadpole, SA, metacercaria, AHC 2829, cysts, 
AHC 2831, 2833 

Cercaria ellisi T.H. Johnston & Simpson, 1944 
Crinia signifera tadpole, SA, metacercaria, 
(experimental), THJ & Simpson 1944: 89 
Tadpole, SA, cyst, AHC 20206 

Cercaria lethargica T.H. Johnston & Muirhead, 

1949 
Tadpole, S A, AHC 2821 

Cercaria natans T.H. Johnston & Muirhead, 

1949 
Limnodynastes tasmaniensis tadpole, SA, 
(experimental), THJ & Muirhead, 1949: 104 
(belongs to Echinostomum group); AHC 12402 

Cercaria sp. 

Tadpole, SA, (K.I. stylet: experimental), AHC 
20260 (Echinostome J: experimental), AHC 



20261 

(Stylet J.W.: experimental), AHC 20262 
Diplostomula 

Hyla aurea, SA, AHC 12390 

Hylaperonii, SA, AHC 12838 

Limnodynastes sp., SA, (experimental), AHC 

12398 

Limnodynastes tasmaniensis, SA, AHC 4125, 

4134, 12702 
Echinostome cysts 

Frog, S A, AHC 12712 

Hyla aurea, SA, AHC 12713 

Tadpole, SA, AHC 12387; (experimental), 

AHC 12722 
Halipegus sp. 

Litoria caerulea, NT, AHC 5405 
Plagiorchid cysts 

Hyla aurea, SA, AHC 12388 
Strigeid cysts 

Hyla aurea, SA, AHC 12384, 12386, 12394 

Limnodynastes tasmaniensis, SA, AHC 12380 
Tetracotyle cysts 

Hyla aurea, SA, AHC 12382 
Digenea cysts 

Bufo marinus, Qld, cysts, Freeland et al. 1986: 

494 

Frog, NSW, cysts, AHC 12393 

Hyla aurea, NSW, cysts, AHC 12372, 12373, 

12390, 12392, 12718-12721 

Hylaperoni, SA, cysts, AHC 12401 

Limnodynastes dorsalis, SA, cysts, AHC 

12369, 12385, 12400, 12406, 12407 
Limnodynastes tasmaniensis, SA, cysts, AHC 

12370, 12371, 12389, 12395, 12397, 12399, 
12406, 12407 

Tadpole, SA, cysts, AHC 12375-12377, 
12403; (experimental), AHC 12379 
Digenea 

Bufo marinus, Qld, Freeland et al. 1986: 496; 

Qld, AHC 15, 19, 2004, 2969, 2971, 2977, 

3145, 3157, 3309, 3313, 3535-3552, 3555- 

3558, 3561, 3562, 3564-3575, 3577-3580, 

3880, 3947, 4077, 4078, 4099, 4101, 4215, 

4351,4889,5020,5021 

Hyla aurea, NSW, AHC 12687, 12681, 4546, 

4537, 4536, 4535; SA, AHC 3520, 4083, 4341 , 

4579, 12688 

Hyla peroni, SA, AHC 12396 

Limnodynastes dorsalis, SA, AHC 3494-3497, 

3499-3501, 3504-3510, 4545, 4548^550, 

12676, 12700 

Limnodynastes fletcheri, SA, AHC 12678 

Limnodynastes sp., SA, AHC 3478-3480, 

3482, 3483 

Limnodynastes tasmaniensis, SA, AHC 1877, 

3484 

Litoria caerulea, Qld, AHC 3522, 3525, 3526, 

12691; NT, AHC 4544 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



19 



Litoria dahlii, NT, AHC 6809, 6993 
Litoria moorei, WA, AHC 8545 
Litoria rothii, Qld, AHC 7181 
Rheobatrachus silus, Qld, AHC 6232 
Taudactylus diurnus, Qld, AHC 8237 

Class Cestoidea Rudolphi, 1808 
Order Pseudophyllidea Cams, 1863 

Family DIPHYLLOBOTHRIIDAE Liihe, 1910 

ILigula sp. 

Hyla aurea, NSW, larval stage, Haswell 1890: 

661 (recorded as having possible affinities with 

Ligula) 

Hyla caerulea, Qld, AHC 2350-2352 

Spirometra erinacei Rudolphi, 1819 
Litoria rubella, NT, AHC 17857 

Diphyllobothriidae spargana 

(IDiphyllobothrium (=Spirometra) erinacei 

(Rudolphi, 1819)) 

Bufo marinus, Qld, AHC 4100 

Hyla aurea, NSW, WA, THJ 1912: 70 

Hyla caerulea, Qld, NSW, THJ 1912: 70 

Hyla latopalmata, ?, (experimental), Sandars 

1953:67 

Hyla latopalmata tadpole, ?, (experimental), 

Sandars 1953: 67 

? Spirometra mansoni (Cobbold, 1882), Stiles & 

Taylor, 1902 Bufo marinus, spargana, Bennett 

1978: 756 

Order Proteocephalidea Mola, 1928 

Family PROTEOCEPHALIDAE La Rue, 191 1 

Ophiotaenia sp. 

Hyla aurea, ?, SJJ 1914: 44; SA, AHC 2825 

Proteocephalus hylae (S.J. Johnston, 1912), 

Prudhoe & Bray, 1982 

syn. Ophiotaenia hylae S.J. Johnston, 1912 
Hyla aurea, NSW, THJ 1912: 63 
Litoria aurea, NSW, QM G 423 
Litoria moorei, WA, BM(NH) 1968.4.19.1-5; 
AHC 8178 

Proteocephalid plerocercoids 

Bufo marinus, Qld, Freeland et al. 1986: 496 
Crinia laevis, Tas, Hickman 1960: 20 
Crinia signifera, Tas, Hickman 1960: 20 
Hyla aurea, Vic, AHC 2327; SA, AHC 8696 
Limnodynastes peronii, Tas, Hickman 1960: 20 

Order Cyclophyllidea Braun, 1900 

Family NEMATOTAENIIDAE Liihe, 1910 

Cylindrotaenia criniae (Hickman, 1960), Jones, 

1987 

syn. Baerietta criniae criniae Hickman, 1960 
Crinia tasmaniensis , Tas, Hickman 1960: 18, 



TMK710-712 

Ranidella tasmaniensis, Tas, Jones 1987: 207 
Cylindrotaenia minor (Hickman, 1960), Jones, 
1987 
syn. Barietta criniae minor Hickman, 1960 

Crinia tasmaniensis, Tas, Hickman 1960: 18 

Crinia laevis, Tas, Hickman 1969: 18 

Crinia signifera, Tas, Hickman 1960: 18; TM 

K716-717 

Ranidella tasmaniensis, Tas, Jones 1987: 21 1 

Assa darlingtoni, NSW, Jones 1987: 212, QM 

GL 4887; Qld, Jones & Delvinquier 1991: 492 

Geocrinia laevis, Tas, Jones 1987: 21 1 

Philoria loveridgei, Qld, Jones & Delvinquier 

1991:492 

Ranidella signifera, Tas, Jones 1987: 211 
Nematotaenia hylae Hickman, 1960 

Hyla ewingii, Tas, Hickman 1960: 8, TM 

K705, K707-709 

Litoria ewingii, Tas, Jones 1987: 184, 185 

Bufo marinus, Qld, Jones & Delvinquier 1991 : 

492 

Crinia signifera, Tas, Hickman 1960: 8, TM 

K706 

Cyclorana novae ho llandiae, Qld, Jones & 

Delvinquier 1991:492 

Limnodynastes ornatus, Qld, Jones & 

Delvinquier 1991: 492 

Litoria fallax, Qld, Jones 1987: 185 

Litoria inermis, Qld, Jones 1987: 185 

Litoria latopalmata, Qld, Jones 1987: 185, 

QM GL 4886 

Litoria pallida, Qld, Jones & Delvinquier 

1991:492 

Litoria peronii, Qld, Jones & Delvinquier 

1991:492 

Ranidella parinsignifera, Qld, Jones 1987: 

185, QMGL 4887 

Ranidella signifera, Tas, Jones 1987: 184, 185 

Ranidella riparia, SA, Jones & Delvinquier 

1991:492 

Uperoleia rugosa, Qld, Jones & Delvinquier 

1991:492 
Nematotaenia sp. 

Hyla caerulea, ?, MRY 1939: 74; NSW, THJ 

1916a: 195, Prudhoe & Bray 1982:12 mf 

Hylafreycineti, ?, MRY 1939: 75; NSW, THJ 

1916a: 1 94, Prudhoe & Bray 1982:12 mf 

Hyperoleia marmorata, ?, MRY 1939: 75; 

NSW, THJ 1916a: 194, Prudhoe & Bray 1982: 

12 mf 
Triplotaenia mirabilis Boas, 1902 

Hyla aurea, ?, MRY 1939: 74 (usually a 

cestode of marsupials; see Prudhoe & Bray 

1982:3 mf for discussion) 

Cestoda Not Further Identified 



20 



DIANE P. BARTON 



Bufo marinus, Qld, AHC 10, 46, 4892 

Crinia signifera, SA, AHC 4419, 4424, 20687 

Crinia sp., SA, AHC 4234 

Hyla aurea, NSW, SJJ 1912: 291; Vic, AHC 

2326; SA, larva, AHC 4584 

Hyla caerulea, NSW, SJJ 1912: 290; Qld, AHC 

1223 

Hyla ewingi, NSW, AHC 4082; SA, AHC 4304, 

4369 

Hyla ewingi alpina, NSW, AHC 4079^081 

Hylafreycineti, NSW, SJJ 1912: 291 

Hyla sp., SA, AHC 40 

Hyperoleia marmorata, NSW, SJJ 1912: 290 

Limnodynastes sp., Qld, AHC 2376; SA, AHC 

2378 

Metacrinia nichollsi, WA, AHC 48 

Rheobatrachus silus, Qld, AHC 8913 

Frog, SA, AHC 20678 



2. Phylum Nematoda 

Class Secernentea 
Order Rhabditida 

Superfamily Rhabditoidea 

Family RHABDIASIDAE Railliet, 1916 

Rhabdias australiensis Moravec & Sey, 1990 
Rana daemeli, Qld, Moravec & Sey 1990: 283, 
CAS N^50 

Rhabdias hylae Johnston & Simpson, 1942 
Hyla aurea, NSW, THJ & Simpson 1942: 176, 
SJJ 1912: 291 (lung nematode); VIC, THJ & 
Simpson 1942: 176; SA, Ballantyne 1971: 51 
Adelotus brevis, Qld, Ballantyne 1971: 51 
Crinia georgiana, WA, Ballantyne 1971: 51 
Crinia glauerti, WA, Ballantyne 1971: 51 
Crinia insignifera, WA, Ballantyne 1971: 51 
Crinia leai, WA, Ballantyne 1971: 51 
Crinia signifera, NSW, SA, Ballantyne 1971: 
50 

Crinia subinsignifera, WA, Ballantyne 1971 : 
51 

Crinia victoriana, Vic, Ballantyne 1971: 50 
Hyla aurea raniformis, Vic, Ballantyne 1971: 
50 

Hyla caerulea, QLD, THJ & Simpson 1942: 
176 

Hyla latopalmata, Qld, Ballantyne 1971: 51 
Hyla lesueuri, Qld, Ballantyne 1971: 51 
Hyla peroni, NSW, SJJ 1912: 290 (lung 
nematode); THJ & Simpson 1942: 178 
Limnodynastes dorsalis, NSW, THJ & 
Simpson 1942: 179 

Limnodynastes fletcheri, Qld, Ballantyne 1971 
51 

Limnodynastes peroni, NSW, SJJ 1912: 290 
(lung nematode); THJ & Simpson 1942: 179; 



Qld, Ballantyne 1971: 51; SA, Ballantyne 

1971:51 

Limnodynastes tasmaniensis , NSW, SJJ 1912: 

290 (lung nematode), THJ & Simpson 1942: 

176; SA, THJ & Simpson 1942: 176, 

Ballantyne 1971: 50; Vic, Ballantyne 1971: 50 

Mixophyes fasciolatus, Qld, Ballantyne 1971: 

51 

Pseudophryne bibronii, NSW, Ballantyne 

1971:50 

Pseudophryne guentheri, WA, Ballantyne 

1971:51 

Pseudophryne occidentalism WA, Ballantyne 

1971:51 

Pseudophryne sp., SA, Ballantyne 1971: 51 
Rhabdias nigrovenosum (Goeze, 1800) 
syn. Rhabdonema nigrovenosum Goeze, 1800; 
listed as a synonym of Rhabdias bufonis 
(Schrank, 1788) in Yamaguti 1961: 84 

Hyla aurea, ?, AM W. 19853-6 
Rhabdias sp. 

Hyla aurea, NSW, VIC, THJ & Simpson 1942: 

178 (referring to 

THJ 1938: 151); WA, BM(NH) 1989.1987- 

1988 

Hyla moorei, WA, BM(NH) 1980.263-282 
Rhabdonema sp. 

Hyla aurea, NSW, Vic, THJ & Simpson 1942: 

178 (referring to Haswell 1891) 

Hyla caerulea, QLD, THJ 19166: 60 

Order Strongylida 

Superfamily Trichostrongyloidea 

Family MOLINEIDAE (Skrjabin & Schulz, 1937) 

Durette-Desset & Chabaud, 1977 

Oswaldocruzia (O.) limnodynastes T.H. 

Johnston & Simpson, 1942 

Limnodynastes dorsalis, SA, THJ & Simpson 

1942: 172; THJ & PMM 1949: 65 

Hyla aurea, NSW, Vic, THJ & Simpson 1942: 

172 

Hyla peroni, SA, THJ & PMM 1949: 65 

Order Oxyurida 

Superfamily Oxyuroidea 

Family PHARYNGODONIDAETravassos, 1919 

Parathelandros australiensis (Johnston & 
Simpson, 1942), Inglis, 1968 
syn. Cosmocerca australiensis Johnston & 
Simpson, 1942 

Limnodynastes dorsalis, SA, THJ & Simpson 

1942: 176 

Limnodynastes fletcheri, S A, Inglis 1968: 173 
Parathelandros carinae Inglis, 1968 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



21 



Heleioporus albopunctatus, WA, Inglis 1968: 

176 

Heleioporus australiacus, WA, Inglis 1968: 

176 

Heleioporus eyrei, WA, Inglis 1968: 176 

Heleioporus psammophilus, WA, Inglis 1968: 

176 

Neobatrachus pelobatoides, WA, Inglis 1968: 

176 
Parathelandros johnstoni Inglis, 1968 

Heleioporus eyrei, WA, Inglis 1968: 175 

Limnodynastes dorsalis, WA, Inglis 1968: 175 

Neobatrachus centralis, WA, Inglis 1968: 175 

(specimens in poor condition, may be P. maini 

or P. limnodynastes) 

Neobatrachus pelobatoides, WA, Inglis 1968: 

175 
Parathelandros limnodynastes (Johnston & 
Mawson, 1942), Inglis, 1968 
syn. Pharyngodon limnodynastes Johnston & 
Mawson, 1942 

Limnodynastes dorsalis, SA, THJ & PMM 

1942: 94; Inglis 1968: 175 

Limnodynastes dorsalis dumerili, SA, THJ & 

PMM 1942:94 
Parathelandros maini Inglis, 1968 

Hyla moorei, WA, Inglis 1968: 176 

Hyla adelaidensis, WA, Inglis 1968: 176 

Hyla cyclorhyncha, WA, Inglis 1968: 176 
Parathelandros mastigurus Baylis, 1930 

Hyla caerulea, Qld, Baylis 1930: 359, Inglis 

1968: 173; NSW, Inglis 1968: 173 

Bufo marinus, Qld, Inglis 1968: 173 

Hyla gracilenta, Qld, Baylis 1930: 359 

Hyla gracilis, Qld, Inglis 1968: 173 (refers to 

Hyla gracilenta recorded by Baylis 1930) 
Parathelandros propinqua (Johnston & 
Simpson, 1942), Inglis, 1968 
syn. Cosmocerca propinqua Johnston & 
Simpson, 1942 

Limnodynastes dorsalis, SA, THJ & Simpson 

1942: 176 
Parathelandros spp. 

Bufo marinus, Qld, Freeland et al. 1986: 496 

Hyla aurea, WA, (female only), BM(NH) 

1980.283-292 

Hyla rubella, WA, (female only), BM(NH) 

1980.318-317 

Oxyurids Not Further Identified 

Bufo marinus, Qld, AHC 2276, 4950; Vic, 

AHC 9048, 9059 

Cyclorqna sp., NT, AHC 4450 

Hyla aurea, Vic, AHC 231 1 

Hyla caerulea, Qld, AHC 2343; NT, AHC 

4947 

Limnodynastes dorsalis, SA, AHC 2306, 3176 

Limnodynastes tasmaniensis, SA, AHC 1417, 

5030 



Litoria rothii, Qld, AHC 7156 
Litoria rubella, Qld, AHC 7180 
Mixophyes sp., Qld, AHC 6172 

Order Ascaridida 

Superfamily Cosmocercoidea 

Family COSMOCERCIDAE (Railliet, 1916 

subfam.) Travassos, 1925 

Cosmocerca limnodynastes Johnston & 
Simpson, 1942 

Limnodynastes dorsalis, SA, THJ & Simpson 

1942: 174 
Cosmocercinae gen. sp. 1 

Rana daemeli, Qld, Moravec & Sey 1990: 273 
Austraplectana kartanum (Johnston & Mawson, 
1941), Baker, 1981 

syn. Rallietnema kartanum Johnston & Mawson, 
1941 

Hylajervisiensis, SA, THJ & PMM 1941 : 146 

Heleioporus eyrei, WA, Inglis 1968: 166 

Hyla moorei, WA, Inglis 1968: 166, BM(NH) 

1967. 1158-1159 

Litoria nasuta, Qld, Baker 1981: 111 

Austraplectana sp. 

Frog, Qld, Baker 1981: 116 
Maxvachonia adamsoni Moravec & Sey, 1990 

Litoria infrafrenata, New Guinea, Moravec & 

Sey 1990: 276, CAS N-449 
Maxvachonia ewersi Mawson, 1972 

Litoria nasuta, New Guinea, PMM 1972: 105 
Maxvachonia flindersi (Johnston & Mawson, 
1941), Mawson, 1972 

syn. Aplectana flindersi Johnston & Mawson, 
1941 ; Austracerca flindersi (Johnston & 
Mawson, 1941) Inglis 1968 

Hylajervisiensis, SA, THJ & PMM 1941: 148 

Bufo marinus, Qld, PMM 1972: 104, AHC 

5170 

Heleioporus australiacus, WA, Inglis 1968: 

165 

Heleioporus barycragus, WA, PMM 1972: 104 

Heleioporus inornatus, WA, PMM 1972: 104, 

AHC 5180 

Heleioporus psammophilis, WA, Inglis 1968: 

165 

Hyla cyclorhyncha, WA, Inglis 1968: 165 

Limnodynastes dorsalis, SA, PMM 1972: 104, 

AHC 5183 

Litoria adelaidensis, WA, PMM 1972: 104, 

AHC 5172 

Litoria caerulea, NT, PMM 1972: 104, AHC 

5182 

Litoria moorei, WA, PMM 1972: 104, AHC 

5175 
Falcaustra hylae (Johnston & Simpson, 1942), 
Chabaud & Golvan, 1957 



22 



DIANE P. BARTON 



syn. Spironoura hylae Johnston & Simpson, 1942 
Hyla aurea, NSW, THJ & Simpson 1942: 173 

Cosmocercoid 

Bufo marinus, Qld, AHC 5009 



Superfamily Ascaridoidea 
Family ASCARIDIDAE Baird, 



853 



Ophidascaris pyrrhus Johnston & Mawson, 1942 

Tadpole, Qld, (experimental infection), QM 

GL9107 

Frog, Qld,QMGZ15 
Raillietascaris varani (Bay lis & Daubney, 1922), 
Sprent, 1985 

Tadpole, ?, QM GL 5674 
Seuratascaris numidica (Seurat, 1917), Sprent, 
1985 

Rana daemeli, Qld, Sprent 1985: 241 

Order Spirurida 

Superfamily Physalopteroidea 

Family PHYSALOPTERIDAE (Railliet, 1893 

subfam.) Leiper, 1908 

Pseudorictularia disparilis (Irwin-Smith, 1922), 

Dollfus & Desportes, 1945 

syn. Rictularia disparilis Irwin-Smith, 1922 

Litoria inermis, Qld, Owen & Moorhouse 

1980: 1014 

Litoria nigrofrenata, Qld, Owen & Moorhouse 

1980: 1014 " 

Rana daemeli, Qld, Owen & Moorhouse 1980: 

1013 
Physaloptera confusa T.H. Johnston & Mawson, 
1942 

Limnodynastes tasmaniensis , NSW, encysted 

larva, THJ & Simpson 1942: 178; SA, encysted 

larva, THJ & PMM 1949:69 

Hyla aurea, NSW, encysted larva, THJ & 

PMM 1942: 91; THJ & Simpson 1942: 178 

Hyla caerulea, Qld, encysted larva, THJ & 

Simpson 1942: 178 

Hyla peroni, SA, encysted larva, THJ & PMM 

1942: 91; THJ & PMM 1949: 69; THJ & 

Simpson 1942: 178 

Limnodynastes dorsalis, SA, encysted larva, 

THJ & PMM 1942: 91; NSW, encysted larva, 

THJ & Simpson 1942: 178 

Limnodynastes dorsalis dumerilii, SA, 

encysted larva, THJ & PMM 1942: 91; THJ & 

Simpson 1942: 178 
Physaloptera sp. 

Cyclorana australis, WA, larva AHC 6399 

Heleioporus eyrei, WA, AHC 3012 

Hyla aurea, SA, AHC 12386 

Limnodynastes dorsalis dumerilii, SA, cysts, 

AHC 2356 (frog taken from intestine of tiger 

snake, Notechis scutatus), 2375 



Superfamily Habronematoidea 
Family HEDRURIDAE Railliet, 1916 

Hedruris hylae Johnston & Mawson, 1941 
Hylajervisiensis, SA, THJ & PMM 1941 : 148 

Hedruris sp. 

Crinia signifera, SA, AHC 28 

Superfamily Filarioidea 
Filarioidea ?gen. ?sp. 

Filaria cochleata Railliet, 1916 
syn. Filaria spiralis Oerley, 1882 

Heleioporus albopunctatus, ?, Oerley 1882: 

312 

Nematoda Not Further Identified 

Agamonema sp. 

Hyla caerulea, Qld, encysted larva, THJ 1914: 
82 

Dorylaimid 

Frog, SA, AHC 6417 

Nematode larvae 

Bufo marinus, Qld, cysts, Freeland et al. 1986: 

496 

Hyla moorei, WA, BM(NH) 1980.298-307 

Arenophryne rotunda, WA, cysts, AHC 6808 

Hyla caerulea, Qld, cysts, AHC 2341 

Nematodes 

Bufo marinus, Qld, Freeland et al. 1986: 496, 
AHC 8,9, 2974, 3258 
Crinia georgiana, WA, AHC 8081, 8079 
Crinia glauerti, WA, AHC 81 19, 81 13 
Crinia haswelli, Vic, AHC 8084 
Crinia leai, WA, AHC 8115, 8082, 8078 
Crinia pseudinsignif era, WA, AHC 8118, 81 14 
Crinia riparia, SA, AHC 8077 
Crinia rosea, WA, AHC 8076 
Crinia signifera, NSW, SJJ 1912:290; SA, 
AHC 20, 22-24, 3617, 6799, 8102, 8105; Vic, 
AHC 1083, 1098; NSW, AHC 8066 
Crinia sp., Vic, AHC 21; S A, AHC 4210, 
421 1, 4214, 4217, 4219, 4231^233 
Crinia subinsignifera, WA, AHC 8080, 8075 
Crinia victoriana, Vic, AHC 8122, 8069, 
8070, 8088, 8096, 8099 
Cyclorana australis, WA, AHC 12880 
Heleioporus eryei, WA, AHC 8120 
Hyla adelaidensis, NSW, AHC 1760 
Hyla aurea, NSW, SJJ 1912: 291, AHC 3528, 
2306, 2308, 2309, 2314-2316, 2318-2321, 
2323, 2324; SA, AHC 3520 
Hyla aurea raniformis, Vic, AHC 8094 
Hyla caerulea, NSW, SJJ 1912: 290, AHC 
2339, 2337, 2336, 2333, 2360; NT, AHC 2331; 
Qld, AHC 2349, 2346, 2344, 2342, 2340, 
2338, 2335, 2235 
Hyla dentata, NSW, SJJ 1912: 291 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



23 



Hyla ewingii, NSW, SJJ 1912: 291; S A, AHC 

8236 

Hylajervisiensis, SA, AHC 1759, 3615 

Hyla lesueurii, NSW, SJJ 1912: 291; Qld, 

AHC 8238 

Hylaperonii, NSW, SJJ 1912: 290; SA, AHC 

12396 

Hyla phyllochroa, NSW, SJJ 1912: 290 

Kyarranus sphagnicolus , NSW, AHC 8247 

Limnodynastes dorsalis, NSW, SJJ 1912: 290, 

AHC 2365, 3362, 

2361, 2360; Vic, AHC 8068; Qld, AHC 2367; 

SA, AHC 2368, 3010, 3176, 8108, 8235 

Limnodynastes fletcheri, NSW, AHC 8059 

Limnodynastes peronii, NSW, SJJ 1912: 290, 

AHC 1728, 3477; SA, AHC 8103 

Limnodynastes sp., Qld, AHC 2605 

Limnodynastes tasmaniensis, NSW, SJJ 1912: 

290, AHC 8064; Vic, AHC 36, 8087, 8100; 

SA, AHC 25, 26, 39, 1877, 1882, 3320, 3619, 

3622, 5031, 8101, 8107, 8110, 12389 

Litoria aurea, SA, AHC 8073 

Litoria booroolongensis, NSW, AHC 8063 

Litoria caerulea, Qld, AHC 8061, 8060 

Litoria dahlii, NT, AHC 6809, 6993 

Litoria ewingii, Vic, AHC 8071, 8072, 8095, 

8097 

Litoria nigrofrenata, Qld, AHC 6145 

Litoria rothii, Qld, AHC 7181 

Litoria verreauxii, NSW, AHC 8085 

Mixophyesfasciolatus, Qld, AHC 8093, 8056 

Neobatrachus pelobatoides, WA, AHC 8121, 

8116 

Neobatrachus pictus, SA, AHC 8104 

Pseudophryne bibronii, Vic, AHC 8090; NSW, 

AHC 8062; SA, AHC 4213, 4218, 4220-1227, 

8089,8106,8111 

Pseudophryne guentheri, WA, AHC 8117, 

8074 

Pseudophryne occidentalis, WA, AHC 8112 

Pseudophryne semimarmorata, SA, AHC 8109 

Uperoleia marmorata, NSW, AHC 8055 

3. Phylum Acanthocephala 

Class Palaeacanthocephala Meyer. 1931 

Order Echinorhynchida Southwell & MacFie, 1925 

Family ECHINORHYNCHIDAE Cobbold, 1876 

Acanthocephalus criniae Snow, 1971 

Crinia tasmaniensis, Tas, Snow 1971: 147, 
TM K228-230, AHC 18165 
Crinia laevis, Tas, Snow 1971: 147 
Crinia signifera, Tas, Snow 1971 : 147 

Pseudoacanthocephalus perthensis Edmonds, 

1971 
Litoria moorei, WA, Edmonds 1971: 55; AHC 



5048,5051 

Limnodynastes dorsalis, W A, Edmonds 1 97 1 : 

55 

Order Polymorphida 

Family PLAGIORHYNCHIDAE Golvan, 1960 

Porrorchis hylae (Johnston, 1914), Schmidt & 
Kuntz, 1967 

syn. Echinorhynchus sp. Johnston, 1912; 
Echinorhynchus hylae Johnston, 1914; 
Echinorhynchus bulbocaudatus Southwell & 
MacFie, 1925; Gordiorhynchus hylae (Johnston, 
1914), Johnston & Edmonds, 1948; 
Pseudoporrorchis hylae (Johnston, 1914), 
Edmonds, 1957 

Limnodynastes dorsalis, SA, encysted larva, 

THJ& Edmonds 1948:69 

Bufo marinus, Qld, encysted larva, Freeland et 

al. 1986: 496 (identified by Edmonds 1989: 

130) 

Hyla aurea, NSW, encysted larva, THJ 1912: 

84, THJ 1914: 83; SA, NSW, THJ & Edmonds 

1948: 69 

Hyla caerulea, Qld, encysted larva, THJ 1914: 

83, THJ & Edmonds 1948:69 

Acanthocephala Not Further Identified 

Acanthocephala sp. 

Hyla caerulea, NSW, QM GL 12287 
Hyla peronii, Qld, QM GL 12346 
Acanthocephala 

Limnodynastes sp., SA, AHC 3409; larva, 
AHC 3481 



Host - parasite checklist 



Order Anura 



Family MYOBATRACHIDAE 

Adelotus brevis (Gunther, 1863) 
N Rhabdias hylae, (lung) 

Arerophryne rotunda Tyler, 1976 
N Nematode larva, cysts 

Assa darlingtoni (Loveridge, 1933) 
C Cylindrotaenia minor, (intestine) 

Crinia georgiana Tschudi, 1838 

N Rhabdias hylae, (lung) 

N Nematodes, (duodenum, rectum) 

Crinia glauerti Loveridge, 1933 

N Rhabdias hylae, (lung) 

N Nematodes, (buccal cavity, rectum, ileum) 

Crinia haswelli Fletcher, 1 894 
see Paracrinia haswelli 



24 



DIANE P. BARTON 



Crinia insignifera Moore, 1954 
N Rhabdias hylae, (lung) 

Crinia laevis Giinther, 1864 
see Geocrinia laevis 

Crinia leai Fletcher, 1898 
see Geocrinia leai 

Crinia parinsignif era Main, 1957 
C Nematotaenia hylae, (intestine) 

Crinia pseudinsignif era Main, 1957 
N Nematodes, (ileum) 

Crinia riparia Littlejohn & Martin, 1965 
C Nematotaenia hylae, (intestine) 
N Nematodes, (rectum) 

Crinia rosea Harrison, 1927 
see Geocrinia rosea 

Crinia signifera (Girard, 1853) 

C proteocephalid plerocercoids, (mesentery & 

under skin) 
C Cylindrotaenia minor, (duodenum, ileum) 
C Nematotaenia hylae, (duodenum) 
C Cestodes, (small intestine) 
N Rhabdias hylae, (lung) 
N Hedruris sp., (stomach) 
N Nematodes, (stomach, intestine, buccal cavity, 

rectum, lung, abdominal cavity) 
A Acanthocephalus criniae, (duodenum, ileum) 
Crinia signifera (Girard, 1853) tadpole 
D Cercaria ellisi, metacercaria, (kidney, 

mesenteries, heart lung), (experimental) 
Crinia subinsignifera Littlejohn, 1957 
N Rhabdias hylae, (lung) 
N Nematodes, (rectum) 

Crinia tasmaniensis (Giinther, 1864) 
C Cylindrotaenia criniae, (duodenum, ileum) 
C Cylindrotaenia minor, (duodenum, ileum) 
A Acanthocephalus criniae, (duodenum, ileum) 

Crinia victoriana Boulenger, 1888 
see Geocrinia victoriana 

Crinia sp. 

C Cestodes, (intestine) 

N Nematodes, (intestine, stomach, rectum) 

Geocrinia laevis (Giinther, 1864) 

C proteocephalid plerocercoids, (mesentery) 

C Cylindrotaenia minor, (duodenum, ileum) 

A Acanthocephalus criniae, (duodenum, ileum) 

Geocrinia leai (Fletcher, 1898) 

N Rhabdias hylae, (lung) 

N Nematodes, (abdominal cavity, duodenum) 

Geocrinia rosea (Harrison, 1927) 

N Nematodes, (rectum) 

Geocrinia victoriana (Boulenger, 1888) 
N Rhabdias hylae, (lung) 
N Nematodes, (duodenum, rectum) 
Heleioporus albopunctatus Gray, 1841 



N Parathelandros carinae, (rectum) 
N Filaria cochleata, (encapsulated between serous 
and muscular layers of stomach) 

Heleioporus australiacus (Shaw & Nodder, 1795) 
N Parathelandros carinae, (rectum) 
N Maxvachonia flindersi, (rectum) 
Heleioporus barycragus Lee, 1967 
N Maxvachonia flindersi 
Heleioporus eyrei (Gray, 1845) 
N Parathelandros carinae, (rectum) 
N Parathelandros johnstoni, (rectum) 
N Austraplectana kartanum, (rectum) 
N Physaloptera sp., (stomach) 
Nematodes, (stomach) 

Heleioporus inornatus (Lee & Main, 1954) 
N Maxvachonia flindersi, (rectum) 

Heleioporus psammophilus (Lee & Main, 1954) 
N Parathelandros carinae, (rectum) 
N Maxvachonia flindersi, (rectum) 

Hyperolia marmorata (Gray, 1 841 ) 
see Uperoleia spp. 

Kyarranus loveridgei (Parker, 1940) 
C Cylindrotaenia minor, (intestine) 

Kyarranus sphagnicolus Moore, 1958 
N Nematodes, (rectum) 

Leptodactylid sp. 

see Myobatrachid sp. 

Limnodynastes dorsalis (Gray, 1841) 

for Limnodynastes dorsalis from any state, except 

WA, see Limnodynastes dumerilii 

N Parathelandros johnstoni, (rectum) 

A Pseudoacanthocephalus perthensis, (intestine) 

Limnodynastes dorsalis dumerilii 

see Limnodynastes dumerilii 

Limnodynastes dumerilii Peters, 1863 

D Gorgodera australiensis 

D Gorgodera sp. 

D Dolichosaccus ischyrus, (intestine) 

D Dolichosaccus trypherus 

D Dolichosaccus sp. 

D Dolichoperoides macalpini, metacercaria, 

(tissues) 
D Digenea cysts 
D Digenea, (intestine, stomach) 
N Rhabdias hylae, (lung) 
N Oswaldocruzia limnodynastes, (intestine) 
N Parathelandros australiensis, (rectum, intestine) 
N Parathelandros limnodynastes 
N Parathelandros propinqua, (rectum, intestine) 
N Oxyurid 

N Cosmocerca limnodynastes 
N Maxvachonia flindersi, (rectum) 
N Physaloptera confusa, encysted larva, 

(mesentery, stomach, peritoneum) 
N Physloptera sp., cysts 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



25 



N Nematodes, (stomach, intestine, rectum) 

A Porrorchis hylae, encysted larva, (mesenteries) 

Limnodynastes fletcheri Boulenger, 1888 

D Dolichosaccus sp. 

D Digenea 

N Rhabdias hylae, (lung) 

N Parathelandros australiensis, (rectum) 

N Nematodes, (duodenum, rectum) 

Limnodynastes ornatus (Gray, 1842) 

C Nematotaenia hylae, (intestine) 

Limnodynastes peronii (Dumeril & Bibron, 1841) 

D Diplodiscus megalochrus, (rectum) 

D Gorgodera australiensis, (bladder) 

D Dolichosaccus anartius, (intestine, rectum) 

D Dolichosaccus trypherus, (duodenum) 

D Haematoleochus australis, (lungs) 

C proteocephalid plerocercoids, (mesentery) 

N Rhabdias hylae, (lungs) 

N Nematodes, (lungs, intestine, rectum, stomach) 

Limnodynastes tasmaniensis Gunther, 1858 

D Diplodiscus microchrus, (rectum) 

D Gorgodera sp. 

D Dolichosaccus trypherus, (intestine) 

D Dolichosaccus sp. 

D Dolichoperoides macalpini, metacercaria 

(tissues) 
D Diplostomula, (buccal cavity) 
D Strigeid, cysts 

D Digenea cysts, (muscles, subcutaneous) 
D Digenea, (gut) 
N Rhabdias hylae, (lung) 
N Oxyurids, (abdominal cavity) 
N Physaloptera confusa, encysted larva, (stomach, 

peritoneum) 
N Nematodes, (lungs, stomach, intestine, rectum) 
Limnodynastes tasmaniensis Gunther, 1858 
tadpole 
D Cercaria angelae, cysts, (wall of thorax and 

rectum, pericardium, tail tissue, base of forleg), 

(experimental) 
D Cercaria natans, (kidney tissue, kidney 

peritoneum), (experimental) 
Limnodynastes tasmaniensis (platycephalus) 
Gunther, 1867 
see Limnodynastes tasmaniensis 

Limnodynastes sp. 

D Diplostomula, (eye), (experimental) 

D Digenea, (stomach, intestine, rectum) 

C Cestodes, (coelom) 

N Nematodes, (stomach) 

A Acanthocephala, (mesentery) 

A Acanthocephala, larva, (rectum) 

Limnodynastes sp. tadpole 

D Cercaria amerianna, diplostomula, (tissues), 

(experimental) 
D Dolichoperoides macalpini, metacercaria, 



(tissues) 
Metacrinia nichollsi (Harrison, 1927) 
C Cestodes 

Mixophyes fasciolatus Gunther, 1864 
N Rhabdias hylae, (lung) 
N Nematodes, (rectum) 
Mixophyes fasciolatus Gunther, 1864 tadpole 
D Fibricola intermedius, metacercaria, (muscles) 

Mixophyes sp. 

N Oxyurid 

Myobatrachid sp. 

D Fibricola intermedius, metacercaria, (muscle) 

Neobatrachus centralis (Parker, 1940) 
N Parathelandros johnstoni, (rectum) 

Neobatrachus pelobatoides (Werner, 1914) 
N Parathelandros carinae, (rectum) 
N Parathelandros johnstoni, (rectum) 
N Nematodes, (rectum) 
Neobatrachus pictus Peters, 1863 
N Nematodes, (rectum) 

Paracrinia haswelli (Fletcher, 1894) 

N Nematodes, (duodenum, rectum) 

Philoria loveridgei Parker, 1940 

see Kyarranus loveridgei 

Pseudophryne bibronii Gunther, 1858 

N Rhabdias hylae, (lung) 

N Nematodes, (duodenum, rectum, stomach) 

Pseudophryne guentheri Boulenger, 1 964 

N Rhabdias hylae, (lung) 

N Nematodes, (rectum) 

Pseudophryne occidentalis Parker, 1940 

N Rhabdias hylae, (lung) 

N Nematodes, (rectum, stomach) 

Pseudophryne semimarmorata Lucas, 1892 

N Nematodes, (rectum) 

Pseudophryne sp. 

N Rhabdias hylae, (lung) 

Ranidella spp. 

for all Ranidella species, see the Crinia equivalent 

Rheobatrachus situs Liem, 1973 

D Digenea, (rectum) 

C Cestodes 

Taudactylus diurnus Straughan & Lee, 1966 

D Digenea, (rectum) 

Uperoleia marmorata Gray, 1841 

for Uperoleia marmorata from all states, except 

WA, see Uperoleia spp. 

Uperoleia rugosa (Andersson, 1916) 

C Nematotaenia hylae, (intestine) 

Uperoleia spp. 

C Nematotaenia sp. 

C Cestodes, (small intestine) 

N Nematodes, (rectum) 



26 



DIANE P. BARTON 



Family HYLIDAE 

Chiroleptes brevipalmatus Peters, 1871 
see Cyclorana brevipes 

Cyclorana australis (Gray, 1842) 

N Physaloptera sp., larva, (buccal cavity) 

N Nematodes 

Cyclorana brevipes (Peters, 1871) 

D Dolichosaccus juvenilis, (intestine) 

Cyclorana cultripes Parker, 1940 

D Allocreadiidae sp. 

D Dolichosaccus juvenilis 

D Mesocoelium microon 

Cyclorana novae hollandiae Steindachner, 1867 

N Nematotaenia hylae, (intestine) 

Cyclorana sp. 

N Oxyurids, (rectum) 

Hyla spp, 

for all Hyla species, see the Litoria equivalent, with 

the following exceptions: 

i) Hyla aurea Lesson, 1829 

for Hyla aurea from NSW (coastal area), see 

Litoria aurea 

for Hyla aurea from SA, Tas, Vic, NSW 

(exclusive of coastal area), see Litoria 

raniformis 

for Hyla aurea from WA, see Litoria spp. 
ii) Hyla ewingi alpina Fry, 1915 

see Litoria verreauxii 
iii) Hyla jervisiensis Dumeril & Bibron, 1841 

for Hyla jervisiensis from all states, except SA, 

see Litoria jervisiensis 

for Hyla jervisiensis from SA see Litoria 

ewingii 

Litoria adelaidensis (Gray, 1841) 

for Litoria adelaidensis from all states, except WA, 

see Litoria spp. 

N Parathelandros maini, (rectum) 

N Maxvachonia flindersi, (intestine) 

Litoria aurea (Lesson, 1829) 

for Litoria aurea from Vic, Tas, SA, NSW 

(exclusive of coastal area), see Litoria raniformis 

for Litoria aurea from WA, see Litoria spp. 

D Diplodiscus megalochrus, (rectum) 

D Diplodiscus sp., (rectum) 

D Distoma sp. 

D Gorgodera australiensis, (bladder) 

D Gorgodera sp., (bladder) 

D Haematoleochus australis, (lungs) 

D Dolichosaccus anartius, (intestine, rectum) 

D Dolichosaccus trypherus, (duodenum) 

D Dolichosaccus sp. 

D Mesocoelium mesembrinum 

D Pleurogenoides solus, (intestine) 

D Digenea cysts, (nerves, muscles, subcutaneous) 

D Digenea, (lung, intestine, rectum) 



C ILigula sp., (muscles, peritoneal cavity, 

subdermal lymph sinuses) 
C Diphyllobothriidae spargana, (thigh muscles) 
C Ophiotaenia sp., (intestine) 
C Proteocephalus hylae 
C Triplotaenia mirabilis 
C Cestodes, (intestine, muscle) 
N Rhabdias hylae, (lung) 
N Rhabdias nigrovenosum, (lung) 
N Rhabdias sp., (lung) 
N Rhabdonema sp. 

N Oswaldocruzia limnodynastes, (intestine) 
N Falcaustra hylae, (intestine) 
N Physaloptera confusa, encysted larva, 

(mesentery) 
N Nematodes, (lung, intestine, rectum, peritoneum, 

abdominal cavity, stomach) 
A Porrorchis hylae, encysted larva, (mesenteries) 
Litoria booroolongensis (Moore, 1961) 
N Nematodes, (rectum, mesentery) 

Litoria caerulea (White, 1790) 

D Diplodiscus megalochrus 

D Diplodiscus sp. 

D Dolichosaccus ischyrus, (intestine) 

D Dolichosaccus symmetrus, (rectum) 

D Dolichosaccus sp. 

D Mesocoelium megaloon, (intestine) 

D Mesocoelium mesembrinum, (intestine, 

duodenum) 
D Mesocoelium microon 
D Mesocoelium sp. 
D Fibricola intermedius, metacercaria, (muscles) 

paratenic host 
D Halipegus sp. 
D Digenea, (intestine) 
C ILigula sp. 

C Diphyllobothriidae spargana, (thigh muscle) 
C Nematotaenia sp. 
C Cestodes, (rectum) 
N Rhabdias hylae, (lung) 
N Rhabdonema sp., (lungs) 
N Parathelandros mastigurus, (small intestine, 

rectum) 
N Oxyurid, (intestine) 
N Maxvachonia flindersi 
N Physaloptera confusa, encysted larva, (stomach, 

peritoneum) 
N Agamonema sp., encysted larva, (stomach wall) 
N Nematode larva, cysts, (intestine) 
N Nematodes, (stomach, intestine, rectum, lung, 

buccal cavity, abdominal cavity, muscle) 
A Porrorchis hylae, encysted larva, (liver) 
A Acanthocephala sp. 
Litoria citropa (Dumeril & Bibron, 1841) 
M Parapolystoma bulliense 
D Mesocoelium oligoon, (duodenum) 
Litoria cyclorhyncha (Boulenger, 1882) 



HELMINTHS OF AUSTRALIAN AMPHIBIA 



27 



N Parathelandros maini, (rectum) 
N Maxvachonia flindersi, (rectum) 

Litoria dahlii (Boulenger, 1896) 
D Digenea 
N Nematodes 

Litoria dentata (Keferstein, 1868) 

N Nematodes, (intestine) 

Litoria ewingii (Dumeril & Bibron, 1841) 

D Diplodiscus microchrus, (rectum) 

D Mesocoelium megaloon, (intestine) 

C Nematotaenia hylae, (duodenum) 

C Cestodes, (small intestine) 

N Austraplectana kartanum 

N Maxvachonia flindersi 

N Hedruris hylae 

N Nematodes, (intestine, rectum, duodenum, 

mesentery) 
Litoria fallax (Peters, 1880) 
C Nematotaenia hylae, (intestine) 

Litoria freycineti Tschudi, 1838 

D Dolichosaccus diamesus, (stomach) 

D Pleurogenoides freycineti, (duodenum) 

C Nematotaenia sp. 

C Cestodes, (duodenum) 

Litoria gracilenta (Peters, 1869) 

D Mesocoelium microon 

N Parathelandros mastigurus, (rectum) 

Litoria inermis (Peters, 1867) 
C Nematotaenia hylae, (intestine) 
N Pseudorictularia disparilis, (stomach) 
Litoria infrafrenata (Giinther, 1867) 
N Maxvachonia adamsoni, (intestine) 

Litoria latopalmata Giinther, 1 867 

C Diphyllobothriidae spargana, (muscles), 

(experimental) 
C Nematotaenia hylae, (intestine) 
N Rhahdias hylae, (lung) 

Litoria latopalmata Giinther, 1 867 tadpole 

D Fibricola intermedins, metacercaria, (muscles) 

C Diphyllobothriidae spargana, (experimental) 

Litoria lesueurii (Dumeril & Bibron, 1 841 ) 
M Parapolystomum bulliense, (bladder) 
N Rhabdias hylae, (lung) 
Nematodes, (rectum) 

Litoria moorei (Copland, 1957) 

D Haematoleochus australis, (lungs) 

D Dolicosaccus trypherus, (intestine) 

D Digenea, (abdominal cavity) 

C Proteocephalus hylae, (intestine) 

N Rhabdias sp. 

N Parathelandros maini, (rectum) 

N Austraplectana kartanum, (rectum) 

N Maxvachonia flindersi, (rectum) 

N Nematode larvae 

A Pseudoacanthocephalus perthensis, (rectum, 



intestine) 

Litoria nasuta (Gray, 1842) 
N Austraplectana kartanum 
N Maxvachonia ewer si 

Litoria nigrofrenata (Giinther, 1867) 

N Pseudorictularia disparilis, (stomach) 

N Nematodes 

Litoria nyakalensis Liem, 1974 

M Parapolystoma sp., (urinary bladder) 

Litoria pallida Davies, Martin & Watson, 1983 
C Nematotaenia hylae, (intestine) 
Litoria pearsoniana Copland, 1961 
M Parapolystoma bulliense, (bladder) 
D Fibricola intermedius, metacercaria, (muscles) 
(natural & experimental) 

Litoria peronii (Tschudi, 1838) 

D Diplostomula 

D Digenea cysts, (rectum) 

D Digenea 

C Nematotaenia hylae, (intestine) 

N Rhabdias hylae, (lung) 

N Oswaldocruzia limnodynastes 

N Physaloptera confusa, encysted larva, 

(mesentery) 
N Nematodes, (lungs, rectum) 
A Acanthocephala sp. 
Litoria phyllochroa (Giinther, 1863) 
M Parapolystoma bulliense, (bladder) 
N Nematodes, (rectum) 
Litoria raniformis (Keferstein, 1867) 
D Gorgodera sp., (bladder) 
D Haematoleochus australis 
D Dolichosaccus trypherus, (intestine) 
D Dolichoperoides macalpini, metacercaria, 

(intestine) 
D Diplostomula 
D Echinostome cysts, (stomach) 
D Plagiorchid cysts 
D Strigeid cysts, (body wall) 
D Tetracotyle cysts 
D Digenea, (intestine) 
C Ophiotaenia sp., (intestine) 
C proteocephalid plerocercoids 
C Cestodes 

C Cestode larva, (abdominal cavity) 
N Rhabdias hylae, (lung) 
N Rhabdias sp., (lung) 
N Rhabdonema sp. 

N Oswaldocruzia limnodynastes, (intestine) 
N Oxyurids, (lung, rectum) 
N Physaloptera sp. 
N Nematodes, (mesentery, intestine, stomach, 

rectum) 
A Porrorchis hylae, encysted larva, (mesentery) 

Litoria rothii (De Vis, 1884) 
D Digenea, (small intestine) 



28 



DIANE P. BARTON 



N Oxyurid 

N Nematodes, (small intestine) 

Litoria rubella (Gray, 1842) 
C Spirometra erinacei 
N Parathelandros spp., (rectum) 
N Oxyurid 

Litoria verreauxii (Dumeril, 1853) 
C Cestodes, (small intestine) 
N Nematodes, (rectum) 

Litoria sp. 

D Dolichosaccus spp. 
D Pleurogenes spp. 
C Cestodes 

Litoria spp. 

identified as Litoria adelaidensis from NSW 
N Nematodes 

Litoria spp. 

identified as Litoria aurea from WA 

C Diphyllobothriidae spargana, (thigh muscle) 

N Parathelandros spp. 

Family RANIDAE 

Rana daemeli (Steindachner, 1868) 

N Rhabdias australiensis, (lung) 

N Cosmocercinae gen. sp. 1 

N Seuratascaris numidica, (stomach, intestine) 

N Pseudorictularia disparilis 

Family BUFONIDAE 

Bufo marinus (Linnaeus, 1758) 

D Diplodiscus sp. 

D Amphistome 

D Dolichosaccus symmetrus, (intestine) 

D Dolichosaccus sp. 

D Mesocoelium mesembrinum, (small intestine) 

D Mesocoelium sp., (intestine, abdominal cavity) 

D Lecithodendriid sp., (intestine) 

D Zeylanurotrema spearei, (urinary bladder) 

D Digenea cysts 

D Digenea, (intestine, stomach, rectum, abdominal 

cavity, lung, buccal cavity) 

C Diphyllobothriidae spargana 

C ISpirometra mansoni, spargana, (muscles) 

C Proteocephalid plerocercoids 

C Nematotaenia hylae, (intestine) 



C Cestodes, (intestine, stomach) 

N Parathelandros mastigurus 

N Parathelandros spp., (intestine) 

N Oxyurid 

N Maxvachonia flinders i, (rectum) 

N Cosmocercoid 

N Nematode cysts 

N Nematodes, (intestine, rectum, abdominal cavity, 

stomach wall) 

A Pororchis hylae, encysted larva 

Unidentified Anura 

Frog 

D Diplodiscus megalochrus, (bladder) 

D Echinostome cysts, (stomach) 

D Digenea cysts 

C Cestodes, (buccal cavity) 

N Austraplectana sp. 

N Ophidascaris phyrrhus 

N Dorylaimid, (intestine) 

Tadpole 

D Dolichoperoides macalpini, metacercaria 

D Cercaria ameriannae, diplostoma 

D Cercaria angelae, cysts, metacercaria 

D Cercaria ellisi, cysts 

D Cercaria lethargica 

D K.I. Stylet cercaria, (experimental) 

D J.W. Stylet metacercaria 

D Echinostome J cercaria, (experimental) 

D Echinostome cysts, (experimental) 

D Digenea cysts 

D Digenea cysts, (experimental) 

N Ophidascaris pyrrhus, (experimental) 

N Rallietascaris varani 



Acknowledgments 

To each of the curators of the parasitic sections in the 
many museums in Australia and overseas that I contacted in 
the preparation of this work I express my deepest gratitude. I 
would also like to thank my fellow PhD students, Mr Steve 
Richards and Ms Sylvie Pichelin, for their much appreciated 
patient assistance with frog and monogenean taxonomy, 
respectively. Drs David Blair, Tom Cribb, Ian Beveridge, and 
Margaret Davies, and Mrs Pat Thomas who offered advice 
and support throughout the preparation of this manuscript, 
also receive my wannest thanks. 



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JOHNSTON, T. H. & MUIRHEAD, N. G. 1949 Larval 
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MATERIAL CULTURE TRADITIONS OF THE WIK PEOPLE, CAPE YORK 

PENINSULA 

P. Sutton 



Summary 

This paper draws upon the author's fieldwork among the Wik people since the 1970s to present a 
summary of the material culture traditions of central-western Cape York Peninsula. Historical 
sources, particularly the ethnographies of the anthropologists Ursula McConnel and Donald 
Thomson, provide additional detail. 



MATERIAL CULTURE TRADITIONS OF THE WIK PEOPLE, CAPE YORK PENINSULA 



P. SUTTON 



SUTTON, P. 1 994. Material culture traditions of the Wik people, Cape York Peninsula. Rec. S.Aust. Mus. 
27(1): 31-52. 

This paper draws upon the author's fieldwork among the Wik people since the 1970s to present a 
summary of the material culture traditions of central-western Cape York Peninsula. Historical sources, 
particularly the ethnographies of the anthropologists Ursula McConnel and Donald Thomson, provide 
additional detail. 

P. Sutton, 7 Edgeware Road, Aldgate, South Australia, 5154. Manuscript received 4 April, 1993. 



Since the 1970s the Australian Aboriginal people 
whose traditions of material culture are presented 
here have been known to anthropologists as 'the 
Wik speaking peoples' or just simply 'the Wik', but 
there is no cover term for them in their own 
languages, nor is there any evidence of such a term 
in the past. They share most of their autogenous 
material culture with their near neighbours, but in 
several ways they remain a recognisable cultural 
entity. Their focal area lies between the Archer and 
Edward Rivers, western Cape York Peninsula, 
Queensland, and inland to Coen. Most Wik people 
still live in this triangle (Fig. 1). 

The Wik include most of the population of 
Aurukun and its outstations (hence 'Aurukun 
people') and that part of Pormpurraaw's population 
known as 'Mungkan side'. They also include west- 
side people affiliated with languages such as 
Mungkanho and Wik-Iiyeyn who live at Coen and 
Meripah in central Cape York Peninsula. The Wik- 
Way, with countries between Aurukun and the 
Mission River to the north, are now also effectively 
Wik people through culture change in the present 
century. Misleading cover-terms for the Wik include 
'the Wik-Mungkan' and 'the Aurukuns', the latter 
sometimes occurring in news media. 

The people discussed in this paper are those who 
are members of clans with traditional estates along 
the coast south from Love River to between 
Christmas Creek and Breakfast Creek, and inland 
from the middle Archer River south via Rokeby and 
Meripah to the upper Holroyd. There are perhaps 
fifty such estates clustered intensively along the 
narrow coastal flood-plain and occupying very much 
larger expanses in the forest and savannah woodland 
country of the uplands. Detailed maps and site 
records for many of these estates are to be found in 
Sutton, Martin, von Sturmer, Cribb & Chase (1990). 
For detailed studies of the traditional land 
relationship systems in the area see especially 
Sutton (1978) and von Sturmer (1978). 



The most dramatic environmental feature of the 
region is the contrast between the food- and 
resource-rich coastal strip, at times only a few 
kilometres wide and subject to massive wet season 
flooding, and the vast, gently undulating hinterland 
with its relatively simple flora and restricted faunal 
range broken only here and there by riverine gallery 
forest and its complex of resources. 

Proceeding inland from the coast, the major 
environmental zones are: 

1. the beach dunes, grading from fine sand to 
coarse shellgrit, sparsely shaded and offering 
limited saltwater resources in the intertidal zone; 

2. the dune woodland along the eastern edge of 
zone 1; extremely rich and complex both florally 
and faunally; 

3. grassy flood plains and huge saltpan/mangrove 
areas, rich in birdlife and, in its freshwater lakes 
or in its tidal estuaries and waterholes, rich in 
marine life; inundated during and after the wet 
season; 

4. another zone of dune woodland on Pleistocene 
sandridges, running approximately in parallel 
with the present coast; rich and complex in 
patches; 

5. relatively open bloodwood and eucalypt forest, 
punctuated by intermittent, broad watercourses 
bearing melaleucas, and containing a large 
number of permanent swamps; 

6. riverine gallery forest fringing permanent 
streams characterised by white sandy beds in 
many sections (chiefly the Archer River system; 
also Kendall River); 

7. grassy savannah woodland with occasional 
outcrops of stone and gravel; this is generally the 
westernmost limit of European pastoral activity, 
although the Aboriginal communities have turned 
off cattle from the coastal area at various times 
since the 1930s. 



32 



P. SUTTON 



Cape Keerweer 



Knox Creek 
Kuchund-eypanh 



CARPENTARIA 



Christmas Creek 




FIGURE 1 . The Wik region. 

SOCIO-CULTURAL GROUPS 

The most important social distinction within the 
Aboriginal population of the region is that between 
'top side' and 'bottom side', a reflection of the 
origins of most of the Wik in countries located either 
inland ('top side') or on the coastal strip ('bottom 
side'). Traditions of material culture also reflect this 
distinction, which is one that has been under- 
recognised by some previous anthropological writers 
(e.g. Thomson 1939a, McConnel 1953). 

Among the coastal peoples it is possible to 
recognise major subgroups consisting of those whose 
clan estates cluster around certain rivers or creeks. 
Thus there are, going from north to south, the Love 
River people, those of the Kirke River ('Cape 
Keerweer people'), Knox River people, Kendall 
River people and so on. These are not linguistic 
groups, but regional groups which have tended to 
closely intermarry, share religious cults, and unite 
at times in conflict with others. In the 1980s and 
1990s the clarity and prominence of these 
distinctions were waning while the top/bottom 



distinction was being strongly maintained. At the 
same time, the old regional ceremonial divisions 
(Apalach, Puch, Wanam etc.) were still known but 
their ceremonies were being performed less often, 
less fully, and in fewer contexts. 

There are no simple political linguistic groups 
('dialectal tribes') in this region (see Sutton 1978, 
Rigsby & Sutton 1980-82). The people do own, by 
right of clan birth and country, a recognised variety 
of languages. In the case of the Wik, all of these 
languages belong to a single genetic family known 
as the Wik group. About fifteen named languages 
constitute this group. This includes languages 
named with the prefix 'Wik-' ['Language-'] (for 
example: Wik-Mungkan, Wik-Iiyanh, Wik-Ngathan, 
Wik-Ngatharr, Wik-Ep, Wik-Me'anh, Wik- 
Keyangan), those named with the prefix 'Kugu-' 
['Language-'] (for example: Kugu-Uwanh, Kugu- 
Muminh, Kugu-Ugbanh, Kugu-Mu'inh), and some 
with no prefix (e.g. Mungkanho). 

Adults and most children have variable 
knowledge of many of these languages, including 
full competence in up to four or five in some cases, 



WIK MATERIAL CULTURE 



33 



although several of these languages were in 
advanced stages of decline and some were virtually 
extinct by the late 1970s (Sutton 1978). By the 
1980s and 1990s Wik-Mungkan and English were 
the dominant languages of the Aurukun area, and 
Wik-Mungkan was the first language of most 
children. Several hundred people, however, had 
partial or excellent knowledge of Wik-Ngathan or 
Wik-Ngatharr, and most languages had at least some 
surviving speakers. 



Prehistory 

The archaeology of the region was virtually 
unknown as late as 1985. Roger Cribb has carried 
out some preliminary survey work (Cribb 1986, 
Cribb et al 1988), paying particular attention to the 
spectacular shell mounds mapped at Love River by 
Cribb, Sutton, Martin and Chase in 1985. The 
Aboriginal custodians of these mound sites, as well 
as their neighbours, consider the mounds to be of 
non-human origin and to be 'story places'. 

Perhaps the most interesting fact about the 
prehistory of the area is that the zone of greatest 
recent population, the coastal plains and dune 
systems, is geologically very young. The coast has 
been prograding, probably under constant human 
occupation, for about 6 000 years (Rhodes 1980). 
The human role in its formation as a mosaic of 
vegetational complexes has yet to be investigated in 
any detail. 



Population 

In 1988 some 900 Wik people lived at Aurukun 
and its handful of outstations. A few hundred more 
lived at Edward River, Coen, Kowanyama, 
Mornington Island and other centres such as Cairns. 
Their numbers had been increasing for some 
decades, after an initial sharp drop caused by 
European, Asian and Islander (Torres Strait and 
Pacific) incursions into the wider region in the late 
nineteenth and early twentieth centuries. 

In 1930 Ursula McConnel estimated the coastal 
Wik people to number two or three hundred (1930: 
99). She thought that 'the Wik-Mungkan' had 
originally numbered 1 500-2 000 people before the 
inroads of exotic diseases (1930: 181). In 1965 the 
Aurukun Aboriginal population, which contained 
most of the Wik people then alive, was 603 (Long 
1970). Only sixteen years earlier the majority of the 
Wik population had been classed as 'nomads', 
living in the reserve but in touch with the mission 
since at least 1938. By 1957, most of the Kendall 
and Holroyd Rivers people had settled at Edward 
River or Aurukun, leaving only a small 
group (Tiger's mob) still living in the bush by the 



1970s (Long 1970: 144-5; J. von Sturmer pers 
comm). 

In pre-mission times population density along the 
coastal plains and dune systems was an estimated 
one person to 2.3 square miles (5.91 square 
kilometres; Sutton 1978: 90). This figure is based 
on a reconstruction using clan maps and an estimate 
of an average 20 persons per clan. A little further to 
the south, in the Mitchell River region, Sharp came 
to a comparable figure of one person to 2.4 square 
miles (6.19 square kilometres) in a census of bush- 
dwelling coastal people in 1932 (Sharp 1940: 487). 
These figures are at the high-density end of the scale 
for Australian Aborigines (see Maddock 1972: 22- 
3) and are vastly higher than what one might expect 
for the hinterland groups, were parallel figures 
available. 



Material culture 



Raw materials 



The coastal Wik region is practically devoid of 
stone and certainly devoid of hard stone suitable for 
tool-making. There are stony ridges, however, in the 
hinterland far to the east. In many months of 
combing the coastal and peri-coastal country while 
mapping old habitation and other sites with Wik 
people this author has only found two stone 
artefacts, both axe heads. Such stone must have 
been traded in from a long distance. Apart from 
local ochres, and the mud and shale used in cooking, 
most raw materials come from plants (wood, bark, 
seedpods, grass stems, leaves, sap) and animals 
(bone, teeth, feathers, shell, spines, wax, hair). 



Settlement and shelter 

In recent decades most Wik people have made 
use of tropical western-style housing when living in 
a settlement village such as that of Aurukun (Fig. 
2). On the fringes of settlements and at outstations 
and temporary bush camps they make use of tin 
sheds, corrugated iron lean-tos, and tents. Leafy 
branches are used for daytime shades and when not 
using a house people would sleep in the open on the 
ground or, in earlier days, on elevated platforms 
during the dry months of the year. Houses and sheds 
were used to store possessions and to keep food 
from the unwanted reach of dogs and visitors, and 
were often too hot to use in the daytime. People 
tended to live under and near, rather than in, a 
modern house, up to the 1980s. 

Thomson (1939a: 218) illustrated and described 
the traditional camps and house types of the region. 
Wet season huts on the coast (Fig. 3) were made of 



VI 



P SUTTON 




FIGURE 2. Modern housing with group preparing for ceremonial house-opening, Aurukun 1987 Photo: Dale Chesson 



melaleuca bark (especially M. leucadendron (L.)L. 
and M. viridiflora). In the inland areas the huts 
were of slringybark (Eucalyptus tetradonia 
Thomson 1939a: 218) or messmate (Garcenia 
warrenii F. Muell. Sutton & Smyth 1980 item 60). 
These two types of hut were given distinct names in 
languages such as Wik-Ngathan and Wik-Mungkan 
and were among the many typifiers of differences 
between the salt-water people and their inland 
neighbours, the fresh-water people. Melaleuca bark 
is the preferred material. 



On the coast, wet season camps were generally 
located under the huge spreading branches of trees 
such as the fig (Ficus microcarpa L.f.) or the 
milkwood (Alstonia actinophylla (A. Cunn.) K. 
S chutn.), where the circular depressions of the 
floors of earlier huts could still be seen in the dune 
sands in the 1980s. These trees were a protection 
against torrential rain and stormy winds. The huts 
were filled with smoke from a small internal fire 
and their one or two small entrances closed off once 
people were inside, to keep mosquitoes at bay. Dry 




FIGURE 3. Wet-season hut of melaleuca bark. Archer River. 1930s. Photo; U. McConnel 



WIK MATERIAL CULTURE 



35 



season day camps also favoured large shade trees, 
and therefore dune woodland or upland forest 
environments, and were often at some distance from 
night camps. A quickly erected day shade was made 
by cutting young saplings and leaning them like a 
brush fence against a pole or spear resting on forks 
of two adjacent trees. A siesta under shade in the 
heat of the day was common practice, especially for 
older people who had spent the small hours of the 
morning awake, tending fires and keeping watch. 

Night bush camps in dry weather were generally 
in entirely open environments such as level parts of 
sand dunes, salt-pans, coastal swales or grass plain. 
The preferred dry season camp was soft sand, free 
of grass or trees, offering long-distance all-round 
visibility as a defence against snakes and human 
intruders, and catching a breeze which would keep 
the mosquitoes down (see Thomson 1939a Plate 
23). 



Fire 



Before matches were introduced, and in remote 
bush camps of the 1970s when matches had run out, 
a fire-drill was used (Fig. 4). Two lengths of young 
wood from the firestick tree (Premna sp.) of 100 to 
150 centimetres were stripped and dried and carried 




FIGURE 4. Fire-making, Cape York Peninsula 1930s 
Photo: U. McConnel. 



in a distinctive bamboo socket bound by yellow 
orchid stem and decorated with jequirity ('crab's 
eyes' or 'gidigidi') beads {Abrus precatorius L.). 
One of the sticks was held on the ground by the foot 
while the other was twirled by a person who stood 
up during the process. While Premna sp. was the 
preferred firestick wood, young stalks of the spear- 
handle tree Hibiscus tiliaceus L. were also used. 

During field work in 1979, Dermot Smyth and 
this author recorded 37 botanically distinct types of 
firewood in the coastal and near inland zones 
between Archer River and Holroyd River. Our 
informants included older people who had lived 
their early lives in the bush, away from the mission. 
Of the various firewoods, 15 were considered good 
to excellent, 19 were considered acceptable, one 
was of use only as a last resort, one was regarded as 
tinder for lighting a fire, and another was only 
mentioned as kindling wood (any dry grass or fine 
dry twigs could be used as kindling). The preferred 
firewoods are marked as such in Sutton & Smyth 
1980. 

Fire was a constant factor in camp life, in travel, 
and in the hunting economy. Decisions on where to 
place hearths, how many hearths were required for a 
camp, how individuals oriented their heads when 
sleeping next to fires, and from whose fire a brand 
could be taken for the starting of another fire, all 
had important implications for the definition and 
negotiation of relationships between individuals and 
groups. In all the Wik languages, 'wife' is 'woman 
fire(-from)' and 'husband* is 'man fire(-from)'. The 
significance of fire and its sharing in traditional 
Western Cape York Peninsula culture is described 
in some detail by Thomson (1932). 

In the course of a day, a particular individual 
might sit at several different fires within the same 
camp. In the daytime, a young bachelor might use a 
fire lit for a brief amount of cooking or boiling a 
billy for tea on a hot exposed patch of sand near a 
day-shade but in the full sun. Back at the base camp, 
in the evening, he might sit and eat with his parents 
or his sister and brother-in-law. For sleep, though, 
he might go to a fire which formed the nucleus of a 
sleeping-place for several other unattached adult 
men. 

When people were dependent entirely on hunting 
and gathering for food, sections of grass-plain were 
fired systematically in the coastal region so that 
small mammals and reptiles could be harvested. As 
the vegetation dried off after the wet season, people 
set fire to the bush wherever they went, in order to 
clear the country for better hunting, for protection 
from snakes, for ease of travel and for improved 
visibility in the constant watch for enemies. These 
fires also marked the positions of bands of people, 
providing important information to their neighbours 
about their direction of travel. In the 1990s it was 



3fi 



P SUTTON 




FIGURE 5. Women digging rush conns, Archer River, 1930s. Photo; U McConnel 




* 
FIGURE 6. Women gathering water-lily seeds and pods. Archer River. 1930s. Photo; U. McConnel, 



WIK MATERIAL CULTURE 



37 



still normal practice to fire country during bush trips 
by vehicle and to light small fires as a signal of 
one's approach to an outstation camp. Rights in the 
firing of one's clan estate were not open to all, 
however, and disputes could arise over wrongful 
firings. 



Subsistence activities 

The basic facts of the seasonal regime and tradi- 
tional subsistence economy in the region are well 
known, and have been the subject of publications by 
Thomson (1939a), McConnel (1953, 1957), and 
Chase and Sutton (1981). (See also von Sturmer 
1978.) The following summary emphasises material 
culture and also contains new information. 

The major vegetable foods in the pre-settlement 
economy were tubers, waterlilies, and an enormous 
variety of fruits. Nuts, the soft inner bark of two fig 
species, and the core of a Livistona palm were also 
eaten. Digging sticks were used, mainly by women, 
to obtain arrowroot {Tacca leontopetaloides (L.) 
Kuntze), round yams {Dioscorea sativa var. 
rotunda) and long yams (Dioscorea transversa R. 
Br.), which constituted staples (Fig. 5). Other tubers 



such as the Cayratia spp. known as kaayketh and 
walken respectively (in Wik-Ngathan) were cooked 
in the ashes (along the coast), or in an earth oven 
(inland), but were not major foods. Of the tubers, 
only long yams were still commonly dug in the 
1980s, as their preparation (brushing the sand off) 
and cooking (in ashes) required little effort. 

Another staple was the lotus lily (Nymphaea lotus 
L. var. australis F. M. Bailey). Its young stems were 
eaten raw and its roots and seeds roasted. Even the 
flowers yielded sweet morsels. It was extremely 
abundant in the wetlands of the coastal region (Fig. 
6). 

Like the arrowroot and round yam, mangrove 
fruits provided abundant foods but also required 
considerable processing. In particular, the grey 
mangrove (Avicennia marina (Forssk.Vierh.)) and 
the small black mangrove (Bruguiera gymnorhiza 
(L. Lam.)) provided bulk carbohydrate, although by 
the 1970s these foods were no longer in any regular 
use. 

The sweeter fruits, mainly growing on jungle 
trees in the dune woodland zones, were usually 
eaten raw, although several species were cooked 
because of their tartness when raw. Sutton and 
Smyth list seventeen fruits of these kinds (1980). 




FIGURE 7. Women using baler shell to obtain water from a well to wash, Archer River, 1930s. Photo: U. McConnel. 



38 



P. SUTTON 






FIGURE 8. Spearing fish with a two-pronged spear and spearthrower, Archer River, 1930s. Photo: U. McConnel. 



The major ones were the yellowfruit or nonda apple 
(Parinari nonda F. MuelL ex. Benth.) which was 
collected in large quantities and stored; the various 
Eugenia and Ficus species (several of each); the 
black cherry {Mallotuspolyadenus F. Muell.); the 
red fruit (Mimusops elengi L.); the wild mango 
(Planchonia careya (F. Muell.) Knuth.); the wongai 
or black cherry (Pouteria sericea (Aiton) Baehni); 
and some of the Terminalia spp.. 

One of the figs, the pandanus and the 'monkey 
nut' tree (Sterculia quadriflda R. Br.) provided nuts 
which required different levels of preparation for 
eating. 

Wild honey from the various Trigona spp. was a 
much sought-after food, occurring in relative 
abundance in the woodland region east of the flood 
plain and on some sandridges in the coastal zone. 
Stone axes (see McConnel 1953: Plate 12: b & p. 24) 
were important for obtaining bee nests from hollow 
trees, but axe heads were traded in from elsewhere 
(McConnel 1953: 11). During the 1970s this author 
found a ground axehead at Uthuk Awuny (Big Lake) 
but it appeared to have had its blade abraded 
through use as a pounder. Hafted stone pounders are 



reported from the area nearby to the south (Thomson 
1936: Plate 8). 

Mice, bandicoots and reptiles such as lizards and 
edible snakes were dug from their nests with 
digging sticks of the same kind used for digging 
tubers. For digging in soft sand or wet mud, baler 
shells were excellent implements and were still in 
frequent use in the 1980s. 

A baler shell on the ground was a common sign 
of the presence of a well. Although potable surface 
water was abundant in the wet season, it was not so 
in the dry season, except inland beyond the 
extensive tidal limits where permanent lagoons or 
the perennial streams of the Archer and Kendall 
Rivers provided fresh water. Coastal people 
preferred to dig a well, even next to a large pool of 
water (Fig. 7). The most preferred water was that 
occurring in the acquifers of ridges based on 
shellgrit. This water is nearest the surface, clears 
quickest after the water level has been dug out, and 
tastes sweetest. Next is sandridge water, followed 
by water which has to be dug in muddy soil. 
Considerable etiquette and religious formality (and, 
occasionally, vehemence) accompanies the digging 



WIK MATERIAL CULTURE 



\9 




FIGURE 9. Morrison Wolmby, Noel Peemuggina and Alan Wolmby standing with spears, Aayk (Cape Keerweeer area) 
1976. Photo. Peter Sutton. 



of wells. Ancestors are informed of who is present, 
requested to do no harm to the diggers, asked to 
make the water come close to the surface, and so 



on. 



Along the coast, fish, sharks and rays were a 
major food source. The actual Gulf beaches were 
not as attractive as the peri-coastal waterways and 
much if not most fishing seems to have been done in 
tidal reaches, estuaries and lakes. A good deal of 
this fishing in earlier times involved the use of 
drives, of weirs or fences across waterbodies, of 
vegetable stupefacients (for example, the bark of 
the bauhinia Cathormion umbellatum (Vahl) 
Kosterm., and Ormosia ornwndii (F. Muell.) 
Merrill), of a variety of nets (ovate, reinforced with 
a cane withy), or of three- or four-pronged spears. 
Rolled melaleuca bark provided a torchlight for 
luring, confusing and illuminating fish during night 
hunting. Women occasionally speared fish but 
spears in general were men's equipment. Multi- 
pronged spears, barbed with bone or, more recently, 
nails, have generally given way to wire-pronged 
spears for obtaining marine species, and shotguns 
and .22 rifles had, by the 1970s, generally replaced 



spears for birds, pigs and wallabies. The old 
wooden-pronged spears were used for spearing birds 
and mammals as well as marine species (Fig. 8), 
and some of the single-pronged spears were used for 
fish, so it is impossible to claim that hunting spears 
were highly specialised. A very common spear type 
was the hardwood-headed spear with a double- 
pointed single barb of bone (later steel). This was 
used mainly for bigger land game but also for 
fishing (see e.g. Thomson 1939a: 210, McConnel 
1953: 26). It is possible, though, to differentiate 
between spears for hunting and those specifically 
made for fighting (see below), even though, in an 
affray, any handy projectile might be used. In the 
1970s in the Cape Keerweer region this author saw 
a stingray 'speared' with a long-handled shovel, fish 
'caught' in a creek with shotgun blasts, and a 
Varanus goanna felled by thrown sticks gathered 
from the ground on the spur of the moment. Similar 
improvisations were probably resorted to in earlier 
times as well. Catching certain marine creatures by 
hand was a prized skill, although some sluggish fish 
in drying pools of water were an easy catch. Water 
birds are said to have been caught by swimmers 



40 



P, SUTTON 




FIGURE 10. Man repairing spear point, Archer River, 1930s. Photo: U. McConnel. 



pulling them underwater. More often the men, 
disguised with mud, and with only their nostrils and 
forehead above water, would swim slowly among 
the birds before startling them into flight. A number 
could then be brought down with a single spear or a 
couple of sticks. 

Large quantities of spears, most of them well over 
two metres in length, were a distinctive mark of a 
western Cape York traditional camp (see Thomson 
1939a Plate 23: lower, and Fig. 9 here). Spear- 
making (and constant repairing) was a major 
occupation for middle-aged and older men (Fig. 10), 
and in the 1980s continued as a source of income. 
The problem of marketing spears through the craft 
outlets was diminished by the introduction of 



detachable sections. Traditional spears were 
sometimes made in three sections, and commonly in 
two, so the adaptation was not a major one. The 
sectioning was designed to yield an optimal 
combination of lightness, strength and flexibility. It 
must be remembered however that, in use, spears 
were essentially fragile and frequently broken or 
damaged. 

Whenever possible, spears were propelled by the 
use of the distinctive Cape YorkPeninsula 
spearthrower with its long, slender body, baler shell 
balance, attached peg, and occasional decorations of 
Abrus seeds and yellow orchid stem binding (Figs. 
8,1 1). A lightweight version capable of floating was 
sometimes used when on the water, and a 'false 



WIK MATERIAL CULTURE 



41 



\ 




FIGURE 1 1 . Spear -thrower made in 1977 by Clive Yunkaporta, Peret Outstation via Aurukun. 



woomera' was used in conflict (see below). 

The inland economy was dependent on fewer 
vegetable staples and a narrower range of other 
vegetable foods, rather widely dispersed. It was also 
focused on freshwater lagoon and river fish and 
shell species and on reptiles. Apart from kangaroos 
and emus, which could not be relied upon as an 
obtainable protein source from day to day, possums 
and bandicoots would have been among the larger 
animals in the diet. By contrast, the coastal economy 
offered not only more variety and greater 
concentrations of foods, but also more abundant 
large-bodied animals such as the wallabies of the 
dune systems, the sharks and rays of the estuaries, 
and, since the 1920s, feral pigs in large numbers. 



Cultivation and domestication 

In pre-mission times people optionally left in situ 
the stem and vine of a tuber they were digging out 
so another would grow on the same vine. In the 
1970s this was still occasionally the practice. By the 
1970s, cultivation of introduced food-bearing plants 
such as watermelon, bananas, pawpaw (papaya), 
cassava and coconuts was also occurring in very 
small plots near outstation camps or planned 
outstation sites, as well as here and there in the 
Aurukun village itself. The ravages of vermin, dogs 
and children, problems with water supplies, and the 
intermittent absence of the garden's personal 
cultivator, frequently led to these gardens falling 
into disuse. Much of their function was to act as 
symbolic claims on place, or as signs of diplomatic 
intent towards Europeans, who have so often 
expressed a desire to see Aborigines interested in 
agriculture. The slightness of their production was 
not critically important. 

As elsewhere in Australia, camp dingoes were 
quickly replaced with European dogs and large 
numbers might become attached to Aboriginal 
households in the area. Dogs continued to be 
regarded as kin and to bear an array of clan-totemic 
names at least well into the 1980s, although most 
also acquired English names. They, like their 
predecessors, were of continuing use as watchdogs 



and of occasional use as hunting aids, and they 
remained kin. 

By the 1970s piglets had become gifts and some 
survived to become members of camps, at least in 
remote places such as Ti Tree. Their tendency to 
bully humans for food can become rather alarming 
by the time they reach full adult size and they 
sometimes meet untimely ends at that stage. 

European cats are occasional pets but their feral 
cousins were not prime hunting targets as they are 
in much of desert Australia. Those kittens which 
survive the attentions of small children might 
develop a loose relationship to certain households 
but do not achieve the valued status of dogs. They 
lack precedent. 

Horses and cattle formed a focus of work and life 
for many Wik people for much of the mid twentieth 
century, although care and maintenance of herds, 
fences, plant and equipment were intermittent and 
depended on a small number of dedicated 
Aboriginal stockmen and occasional European 
managers. While the strong arm of the mission was 
in charge, the Aurukun cattle industry turned off 
beasts, barging them out off the coast or walking 
them overland as far as the railhead at Mungana 
some 400 kilometres to the southeast. In the 1980s, 
after the cattle operations had been in disarray and 
decay for a decade or more, Aurukun Community 
Incorporated organised the eradication of cattle from 
the Aurukun Shire as part of the Australia-wide 
brucellosis and tuberculosis eradication campaign. 



Food preparation and consumption 

While heavy game might be transported tied to a 
long pole carried by two people, under bush 
conditions most foods were carried in woven bags 
or in containers made from hardwood bark (Fig. 
12). Food was also wrapped in paperbark (from 
Melaleuca spp.) and tied with vines or grass, both 
for transportation and warming near a fire. 
Paperbark is in fact the commonest and most 
versatile natural material in the western Cape York 
traditional subsistence kit (Figs. 13,14). The uses of 
paperbark extend well beyond subsistence, but 



42 



P SUTTON 




FIGURE 12. Women with heavy loads, Archer River, 1930s. Photo: U. McConnel. 



because of its special role in cooking and eating we 
will deal with all of its uses here. 

Paperbark is used as a food preparation surface 
on which meat can be sliced or a damper kneaded 
(Fig. 15), a coverlet to shield food from flies or 
dogs, a source of kindling when outer barks and 
grasses are wet, a heat-sealing medium for meats in 
earth ovens, a mitt with which to pick up steaming 
hot foods or billycan handles, a platter from which 
to eat, a towel with which things are wiped, a lightly 



abrasive cleaner (when crumbled) for greasy or 
blood-covered hands, a clean, dry surface to sit or 
sleep on, a blanket to sleep under, a shroud in which 
to mummify and transport corpses, a roofing 
material for shelters, a torch for night travel or 
fishing, a pouch for containing stingray barbs or 
small medicinal or magical substances, a cigarette 
paper, a binder for awl handles, a napkin for babies, 
a menstrual pad, and a sheet of toilet paper. Its 
abundance in coastal swamp areas is one of the 



WIK MATERIAL CULTURE 



43 




. * ,' . 



FIGURE 13 
Archer River. 



- ^F-^^-w^^mm. 



■ :^'r** ■. . ■* 



Stripping paperbark from a melaleuca tree, 
1930s. Photo: U. McConnel. 



factors which makes the coastal region such a 
convenient place to live, compared with the inland, 
The preferred species provide sheets of soft, durable 
material which arc easily prised from the tree with a 
sharp stick, which impart little or no odour or taste 
to food and which leave no sharp or stringy 
fragments behind when fragmenting. 

Most cooking was carried out by simple broiling 
on an open fire, or by the earth oven technique. 
Shark or ray liver, however, might be fried in a 
baler shell, or lightly cooked before being wrapped 
in the washed white flesh of the shark or ray and 
then tied up in paperbark to be slowly warmed 
through at the hearlhside. This distinctive loaf of 
high vitamin and protein content gives rise to the 
term by which most Wik people refer to the 
elasmobranch (sharks and rays) category: Hying 
meats' (for example, minh katheng in Wik- 
Ngathan). The sign for the same category is the 
wringing of hands, a reference to the squeezing of 
white fluid from the flesh after cooking and before 
eating. 

The earth oven technique was used for large 
game, but also for large quantities of small game 
such as marsupial mice and fruit bats or vegetable 
foods such as tubers. The following description of 
that method is probably only strictly true for the 
coastal region, where there is no stone. (Note also 
that other cooking practices varied within the region: 
inlanders cooked the tuber known as angk (Wik- 
Mungkan) or walken (Wik-Ngathan), probably a 
Cayratia sp., in the earth oven, but coastal people 
roasted it in the coals; some coastal clans half- 
cooked stingray livers, while others placed them raw 
inside the white flesh. These differences were 




FIGURE 14. Carrying sheets of paperbark. Archer River. 1930s- Photo: U. McConnel. 



44 



P. SUTTON 




FIGURE 15. Paddy Yantumba removing eggs from a file 
snake. Big Lake 1976. Photo: Peter Sutton. 



consciously maintained as an aspect of local group 
identity.) 

To make an earth oven, a pit was dug and a 
vigorous blaze lit in it (Fig. 16). On this fire, or one 
nearby, the fur, skin or scales of the beast were 
singed away and the burnt fragments lightly scraped 
off. Lumps of termite mound, if available, were 
thrown into the pit to get hot. (The earth oven 
technique is known as '(burying) in termite 
mound'.) Where there was no termite mound - 
whose heat-holding properties are exceptional - 
lumps of shellgrit were used. At the bottom of this 
hierarchy of oven bricks was swamp mud, a last 
resort when the other two substances could not be 
had. Termite mounds arc thus a factor in defining 
optimal campsites in the region. 

When the oven bricks were hot, they were 
covered in green leaves on which the meat was 
placed skin-side up. Large sheets of paperbark (or, 
in the 1980s, corrugated iron if available) were then 
placed on the meat and the whole oven was sealed 
with sand. The relative ease with which the pit 
could be dug and the oven properly sealed depended 
on the presence of sand, and the whole process 
usually required access to suitable paperbark trees, 
which were thus further factors in defining a good 
campsite location. Earth ovens were usually near, 
not right on, overnight and base camps. They 
accumulated quantities of offal, skin, feathers and 
bones which made the immediate area unpleasant 
for camping. The termite mound lumps were re- 
used until fragmented, on subsequent visits to the 
site. 

Limited food storage was practised. Nonda plums 




FIGURE 16. Earth oven cooking, Watha-nhiin Outstation, 1976. Photo; Peter Sutton. 



WIK MATERIAL CULTURE 



45 



(Parinari nonda F. Muell. ex. Benth.) were dried 
on the rooves of shelters, or collected dry from the 
ground (the dry form even has a different name), 
and were kept for some weeks after their season of 
superabundance. Long yams (Dioscorea transversa 
R. Br.) were stored in the sand for weeks and even 
months (Sutton and Smyth 1980). Long-necked 
turtles might survive a day or two trussed up, and in 
the Big Lake area barramundi is said to have been 
cooked, wrapped in paperbark, and buried in the 
cool earth for eating days later. Most food, however, 
was eaten within twenty four hours. 

Food preparation was elaborate in the case of 
vegetable foods with toxic or unpleasant properties 
which required scraping, pulverising, leaching and 
sieving (for example, the round yam Dioscorea 
saliva, the arrowroot Tacca leotopetaloides (L.) 
Kuntze, or the mangrove fruit Avicennia marina 
(Forssk.) Vierh.). These were not preferred foods 
and by the 1970s most had become just a hardship 
memory. Nonda plums were pounded and mixed 
with water to make a kind of fruit soup (Thomson 
1939a), much favoured by children and by the 
toothless. A wooden mallet and pounding board 
were used in these processes, and mashed food was 
collected in large messmate bark containers (Fig. 
12). This same bark (from Garcinia warrenii F. 
Muell.) was also used for canoes, sleeping 
platforms, and inland shelters. It is a characteristic 



resource of the hinterland behind the narrow coastal 
floodplain. 

Foods were mixed in earth ovens in order to 
create changes of flavour, and some non-foods were 
used as condiments in the same context. For 
example, the leaves of two eucalypts known in 
English as 'bloodwoods' (but named separately in 
Wik languages) were placed in earth ovens with the 
meat of game such as wallaby or wild pig to 
improve their flavour. This attention to culinary 
detail marks the culture of this kind of region as 
very different from, for example, the cooking styles 
of people of the Western Desert. 



Travel and transport 

By further contrast with inland peoples, and 
especially those of desert Australia, in pre- 
settlement times Wik people were comparatively 
sedentary, making less frequent shifts of camp and 
travelling much shorter distances between camps. 
Base camp shifts were usually about five or six 
kilometres only. People were therefore able to carry 
more possessions, and lived in an environment rich 
in a wide variety of raw materials from which 
artefacts might be made. It is not surprising, then, 
that the inventory of their material culture is 
relatively large. 




FIGURE 17. Noel Peemuggina and Johnny Ampeybegan drinking from baler shells at the beach near Big Lake, 1976. 
Photo: Peter Sutton. 



46 



P. SUTTON 




FIGURE 18. Woman weaving basket, Archer River, 1930s. 
Photo: U. McConnel. 



Message sticks, which were small tablets of wood 
carved with non-figurative symbols representing 
days of travel, places, people or commodities, were 
carried by messengers when arranging meetings or 
other dealings, both ceremonial and secular 
(McConnel 1953, Sutton 1978: 93-4). 

The variety of containers used to transport or hold 
things in this region was, perhaps, exceptional. In 
addition to the woven bags and baskets discussed 
below, and the bark containers discussed above, 
there were the ubiquitous baler shell (used for 
drinking (Fig. 17), digging, baling and cooking as 
well), the conch shell and bamboo tube (for 
transporting water), the palmleaf cup, and the 



carved wooden vessel (see Thomson 1939a: Pl.XXI, 
1939b: 85). 

In the 1980s women of the Wik peoples were still 
making a variety of differently shaped woven bags 
and baskets employing many different fibres and 
weaves, and a number of natural dyes (see Adams 
1986, McConnel 1953, Thomson 1939a). These 
were used not only for carrying things, but also for 
sieving and leaching bitter foods, for hanging 
valuables high above the reach of children and dogs, 
for imparting a militant spirit to small boys (by 
smacking woven bags against their calves), and, in 
the case of larger woven baskets, as cradles for 
babies. Woven containers were also a focus of 
traditional religious symbolism (there was a Woven 
Bag totemic clan, for example) and, because of their 
frequent use as gifts, they were an important 
medium for maintaining good relations between 
individuals and groups (Figs. 18,19). 

The two main types of watercraft were the simple 
'floating log', most frequently used during brief 
river crossings, and the messmate bark canoe. From 
the Archer River north, dugout canoes were made 
from the cotton tree and used for hunting sea turtle 
and dugong in the Gulf waters. Canoes were used 
on the inland lakes, swamps and estuaries during 
and just after the wet season, particularly for 
collecting eggs of the magpie goose, which are 
superabundant at that time, and for spearing fish. 
They were both paddled and punted. Figure 20 
shows a canoe made and used for egg collecting in 
the Munpun area in the wet season of 1975-6, and 
then abandoned. That may have been one of the last 
traditional uses of a canoe in the region, as 
aluminium dinghies have become commonplace and 
store foods have increasingly replaced bush foods. 




FIGURE 19. Basket made by Isohel Wolmby from Aurukun. Collected by P. Sutton, 1986 



WIK MATERIAL CULTURE 



47 





id'- ^ 




:.Mjjjsa' 



.■■■■■,.. ■!■; 






i:K.^3K' : ' ^WF: : : •S? IIP ■ ^IP- 

figure 20. Disused bark canoe near goose-egg swamps, Munpun 1976. Photo: Peter Sutton. 



Fighting and duelling 

Women fought with yardsticks and men fought 
with spears and spcarthrowers, particularly if the 
conflict took place among those who had had time 
to prepare for this important aspect of individual 
and group relationships in the region. A 'false 
woomera' was also used by men - this was a 
spearthrower lacking a peg or baler shell ornament 
or both - and was used as a club. Spears for fighting 
at a distance were shorter and unbarbed. Long 
fighting spears were used for closer combat and for 
jabbing in the thigh as a means of settling 
grievances (see McConnel 1953: 25-6). Some of the 
spears specifically made for fighting had a cluster of 
stingray barbs at the nose, all pointing forward. 



These made an extremely painful and messy wound 
(Fig. 21). Men's clubs were either long and pointed, 
like yamsticks, or short, knobbed and pointed, and 
were used both for impact and for fending off spears 
(Fig. 22). The long throwing stick was also used to 
deflect spears. Shields were not used in this area. 
The proboscis of a sawfish or the teeth of a shark 
would be set in a handle of milkwood {Alstonia 
actinophylla (A. Cunn. A. Schum.)) to form a 
fighting sword. The conflict-related part of the 
material culture array in this region was clearly 
highly elaborated. 

By the 1980s conflict was carried on primarily 
with fist-fighting and the opportunistic use of 
objects at hand which might be turned into weapons, 
such as pieces of fence, billy cans, bottles. 













iff :J:::-''-^'l--.;--:' ■'' 
™:t :: ir.V ■■ 


-TT^^^^^^p 



































FIGURE 21 . Spear-head showing sling-ray barbs, made in 1977 by a Wik man, Peret Outstation. 



48 



P. SUTTON 




lllli;^;:;-:;; 




FIGURE 22. Two-pronged wooden club, decorated with seeds traded from Torres Strait, used to parry spears, Archer 
River, 1930s. Collected by U. McConnel. 



broomsticks, steel knives and, occasionally, rifles or 
shotguns. By the early 1990s spearings had become 
rare. 



Manufacturing technology 

Tools used in manufacturing traditional artefacts 
included knives (originally made from shell, 
especially mudshell, or from bamboo, but latterly of 
steel), shell drills for boring holes in necklet shells, 
axes made of stone or steel, paired sticks used to tie 
the two sides of bags during weaving, bone awls 
used to pierce bark for canoe-making, wallaby 
incisors (still in the jaw) for cutting and graving, 
and a 'palette' or resin bat made of ironwood, with 
a wallaby incisor set in its handle, which was used 
for smoothing heated resin or wax and for engraving 
(Fig. 23). Woodrasps, saws, hammers and nails, 
heated wires, sharpening stones and butcher's 



'steels' were all in regular use by a cross-section of 
Wik peoples by the 1970s. Carpentry and basic 
mechanics' skills were taught by missionaries and 
other community workers and a wide range of 
workshop tools were available in the main 
population centres from about the 1930s. 



Magic and medicine 

As in many parts of Aboriginal Australia, 
ironwood smoke (using the leaves of Erythrophleum 
chlorostachys (F. Muell. Baillon) in this case) was 
used to send away the spirits of the dead. After 
scraping, the roots of a fern (Drynaria quercifolia 
(L.) John Smith) were burned to yield a smoke 
which would send people into a deep sleep so one 
could, for example, steal away from one's family at 
night to engage in sexual activity. One could make 
oneself invisible (for example, when seeking 




FIGURE 23. Ironwood palette used for smoothing heated wax and as an engraving tool, with wallaby tooth incisor 
attached. Collected by U. McConnel, Archer River, 1930s. 



WIK MATERIAL CULTURE 



49 



revenge) by tying one's upper arms with a string 
made from the roots of a fig (Ficus virens Aiton ex 
Dryander) or with the grass called keent in Wik- 
Mungkan (Family Gramineae), Rain could be 
prevented by burning the leaves of the whitefruit 
[Eugenia eucalyptoides F.Muell.), or by burning the 
roots and branches of the deciduous hardwood 
known in Wik-Mungkan as yuk kuk (possibly 
Tinospora minasira). Gamblers could improve their 
luck by secreting on themselves pieces of the bark 
of Excoecaria parviflora Muell. Arg., or of the 
'crocodile-hand tree' {Terminalia subacroptera 
Domin.). These and other magical substances come 
under the same cover-term as materials used for 
both healing and sorcery in each of the Wik 
languages (for example, operr, Wik-Mungkan, and 
'medicine' or 'bush-keymas [chemist]' in English, 
although the Pacific pidgin loan purriy-purriy is 
used for sorcery items). 

Magical procedures in this region were complex, 
and too many to give in full here. One example is 
the transformation of wooden effigies or real 
specimens of small reptiles into live saltwater 
crocodiles which were then owned and controlled 
(and considered 'sons') by particular older men. The 
effigy or lizard would be bound with string and 
smeared with blood - preferably human - before 
being released into the water with appropriate 
exhortations. On returning to camp, the crocodile 
man {pom thikel-kathenh, 'man crocodile-ties' in 
Wik-Ngathan) must bring home very large pieces of 
firewood, not small ones, otherwise the crocodile 
would be undersized. These men are said to have 
called up their personal crocodiles and to have stood 
in the water with them, cleaning their teeth with a 
twig. The crocodiles could be sent to the river of 
another group to kill people. Deaths from crocodile 
were normally attributed to the malevolence of 
others. 



Ceremonial life 

Ceremonies in the area involve elaborate carvings 
made of wood, hair, bone and many other materials 
(see McCarthy 1964, 1978; Berndt, Berndt & 
Stanton 1981; Morphy 1981; Sutton 1988; Bartlett 
1989). These are among the most spectacular 
sculptures in Aboriginal Australia. 

While they are highly traditional in meaning, and 
carved and painted totemic wooden objects were 
collected from the region at the earliest stages of 
contact, in their present elaborate form the Aurukun 
sculptures appear to have flourished mainly since 
the advent of steel tools and the introduction of 
techniques such as morticing during the mission 
period in the present century (see McCarthy 1964: 
300). These more technically complex works 



probably do not pre-date the late 1940s, when a 
major collection of them was made at Aurukun and 
lodged with the University of Queensland (now in 
the Anthropology Museum, Department of 
Anthropology and Sociology). The many examples 
filmed in ceremonial use at Aurukun by the 
Australian Institute of Aboriginal Studies in 1962, 
and collected by Fred McCarthy, are now in the 
National Museum of Australia (Dunlop 1964, 
McCarthy 1978). Although they were still being 
made for ceremonies in the 1990s, and some had 
been given to outside individuals or institutions 
(such as the South Australian Museum), only rare 
examples were allowed to reach the market. 

These sculptures are used in regional cult 
ceremonies, a highly competitive form of religious 
celebration in which events from local mythologies 
are enacted by dancers to the accompaniment of 
songs and symbolic cries. They are, among other 
things, statements about rights and interests in 
specific places. They refer to the mythic 'title deeds' 
of Aboriginal customary law, and their making and 
use in performance by particular people wearing 
particular body-paint designs have strong local 
political and territorial meanings, as well as spiritual 
and aesthetic aspects. They may be spiritually 
dangerous for some time after their manufacture and 
are usually allowed to rot away in the bush. 

Most Aurukun sculptures are figurative 
representations of particular beings in the myths, 
and are generally long and gracile, ranging from 
around 400 mm to over two metres in length 
(published illustrations are in e.g. McCarthy 1964, 
Morphy 1981, Berndt, Berndt & Stanton 1982, 
Sutton 1988). They are painted in non-realistic 
bands and patches of colour, and show a distinctive 
degree of trouble taken to insert realistic (and 
sometimes real) teeth, spines, tails and fins into the 
figures. They lack the smooth, static formalism and 
inscrutable stolidity of a good deal of the carving 
associated with some other so-called tribal societies. 
They have a stark, unpredictable and dramatic look 
which appeals especially to lovers of modernist 
'primitivism'. 

More restricted ceremonial material cannot be 
reported on here (cf. McConnel 1953), but it is 
appropriate to note that, according to initiated men 
alive in the 1970s, a previously unreported drone 
tube was in regular use at initiation ceremonies up 
to the late 1960s, although none have been collected 
to the author's knowledge. Percussion 
accompaniment to singing was mainly done by 
clapping. Saliva was regularly licked onto the hands 
to increase the clapping volume. Performers wore 
iridescent pendants cut from mother of pearl or 
nautilus shells, which were smeared with red ochre. 
They also wore filaments of white cockatoo feathers 
joined at the stem with a resin/wax mixture, 



so 



P. SUTTON 



armbands of red-ochred pandanus, and sometimes 
armbands and girdles made from bark studded with 
red Abrus seeds. Some of these ornaments were also 
used in daily life. 

When performing 'Island Dance', a dancing and 
singing style based on an amalgam of Pacific Island 
traditions with those of Cape York Peninsula, 
dancers in recent decades have worn and carried 
matchbox bean rattles, and sported colourful cotton 
nagas (loincloths) or synthetic grass skirts. Rhythm 
for this style is provided by hand-drumming on a 
membrane stretched over the end of a wide cylinder 
(often an inner tyre tube on a length of plumber's 
polythene pipe), or by beating of sticks on a metal 
flour drum. In the earlier part of this century, 
cylindrical drums were only used north of Archer 
River (McConnel 1953: 23). By the 1980s women 
had added a very erotic version of Polynesian hula 
dancing to their repertoire, which required flowers 
in the hair, leis, and brightly coloured synthetic 
grass skirts (Lurex is desirable). But hula here is 
only performed at mortuary ceremonies. 



Personal adornment 

Before adopting western dress, Wik people 
basically went naked, although women wore string 
aprons for symbolic or ritual reasons at certain times 
(see McConnel 1953: 15), People did wear many 
ornaments though, including pendants of shell and 



,^m 





FIGURE 24. Widow's necklet, with beeswax pendants 
studded with Abrus seeds. Collected by U. McConnel, Cape 
York Peninsula, 1930s. 



■ ■ . ' ■: ■ 



FIGURE 25. Shell pendant made by George Sydney 
Yunkaporta, 1977, Watha-Nhin Outstation. Collected by P. 
Sutton. 

beeswax, shell nosepegs and wooden earlobe 
cylinders, strings and girdles of hair, fur, feather 
down, orchid-bark, palm fibre string, fig 
pneumatophore string, and threaded cowrie shells, 
pearlshell pieces, grass-bugles and Abrus seeds 
(Figs. 24,25). These adornments reflect an aesthetic 
preference for white and shining objects, or red 
objects. White clay and red ochre are also the 
dominant colours of ceremonial body paints in the 
region. 

Aboriginal footwear is usually associated mainly 
with desert groups, but this author has been told on 
reliable authority that Wik people in the past did 
make grass and string sandals. Anyone who has 
tried to walk on the region's hot dry sands in bare 
feet will understand why this may well have been 
so. Local languages have an autogenous term for 
footwear (tha' morrok) which may have originally 
referred to the sandals but now refers to shoes and 
boots. 



Drugs 

Thomson (1939b) came to the view that tobacco 
had been available, but not grown, in Cape York 
Peninsula for a very long period before his first visit 
there in 1928. He reported three kinds of smoking 
pipe. One was a long cylinder, usually of bamboo or 
ironwood, open at one end into which someone 
smoking a cigarette or pipe expelled smoke which 
was at the same time inhaled by someone else 
through a small hole near the other end. Another 
was a short, broad cylinder of bamboo or ironwood 
which was filled with smoke then passed around to 
others who inhaled and exhaled the same smoke 



WIK MATERIAL CULTURE 



5) 




■ 




FIGURE 26. Communal smoking-pipe, collected by U. McConnel, Archer River, 1930s. 



(Fig. 26). The third was modelled after the English 
briar-pipe, made of a very hard species of wood, and 
bored out with a hot wire. This last type was the 
only one still in regular use among the Wik peoples 
by the 1970s, although cylinder pipes were still 
made for the Aurukun craft shop. 



Material culture today 

At the present time most Wik people live for most 
of the year in modern Queensland-style houses or 
the tin sheds of outstations, and move about by 
Toyotas, cars, planes and tractors or motor-powered 
boats and, occasionally, horseback. Television sets 
and videos have become common possessions. Some 
Aboriginal households have acquired telephones. 
While traditional ceremonies are maintained in an 
attenuated form, Island Dance, hula and rock'n roll 
have become more frequently performed. The dead 
are no longer mummified, carried about for long 
periods or cremated at elaborate ceremonies, but are 
buried with a simple Christian fite and their houses 
ritually 'opened' by a mixture of traditional totemic 
and modern secular performances. Their spirits are 
still ritually sent to their homelands and their names 
are still prohibited from public use for a period after 
the death. 

By the 1980s, the Wik economy had become 
based largely on welfare payments and limited local 
employment on community services, although fish, 
pigs, crustaceans and water birds still provided 
significant proportions of the diet. Alcohol had 



become a major economic and social preoccupation. 
Rifles, shotguns, nylon fishing lines and steel hooks 
had replaced most kinds of traditional hunting 
equipment, although the multi-pronged wire spear 
and spearthrower were still used for shoreline 
hunting. A number of bush medicines were still 
used, but constant use was also made of western 
medicines obtained through community hospitals 
and health workers. Many traditional items of 
material culture, and some objects introduced by 
missionaries (such as feather flowers and pandanus 
place-mats and bowls), were still made, however, 
either for the cash available from the craft outlets 
(see Adams 1986), or for gifts within the 
community. 



Acknowledgments 

My chief debt is to the people of the Cape Keerweer region, 
especially the residents of Watha-nhiin and Aayk Outstations 
in the years 1976-1979. Noel Peemuggina, Isobel Wolmby, 
Ray Wolmby, Clive Yunkaporta and Peter Peemuggina were 
my main informants on the specific matters dealt with here. I 
also wish to thank Derrnot Smyth for making the botanical 
collections and for his cheerful company over many weeks of 
arduous field work. Derrnot Smyth and The Queensland 
Herbarium provided the botanical identifications. I thank the 
South Australian Museum for permission to reproduce 
Figures 3-8, 10, 12-14, and 18, and the Museum's 
photographer, Trevor Peters, for Figures 11, 19, and 21-26.. 
Funding for field work was provided by The Australian 
Institute of Aboriginal and Torres Strait Islander Studies, the 
Australian Department of Education, and the University of 
Queensland. 



References 



ADAMS, J. 1986. Crafts from Aurukun. Design for a local 
environment. Aurukun. Queensland: Aurukun Community 

Incorporated. 

BARTLETT, J. 1989. Australian anthropology. Pp. 13-70 in 
'Cultural exhibition of Queensland'. Ed. J. Bartlett & M. 
Whitmore, Saitama Prefectural Museum: Saitama. 

BERNDT, R. M ., BERNDT, C. H. & STANTON J. 1981. 
'Aboriginal Australian art'. Methuen: Sydney. 



CHASE, A. K. & SUTTON, P. 1981. Hunter-gatherers in a 
rich environment: Aboriginal coastal exploitation in Cape 
York Peninsula. Pp.1 8 17-1852 in 'Ecological biogeo- 
graphy in Australia'. Ed. A. Keast. W. Junk: The Hague. 

CRIBB, R. 1986. Archaeology in the Aurukun region: a 
report on work carried out in 1985. Queensland 
Archaeological Research 3: 133-58. 

CRIBB, R., WALBENG, R., WOLMBY, R. & Taisman, C. 



52 



P. SUTTON 



1988. Landscape as a cultural artefact: shell mounds and 
plants in Aurukun, Cape York Peninsula. Australian 
Aboriginal Studies 1988(2): 60-73. 

DUNLOP, I. (director) 1964, Dances at Aurukun. Australian 
Commonwealth Film Unit for Australian Institute of 
Aboriginal Studies: Sydney. Colour film, 16 mm, 28 mins. 

LONG, J. P. M. 1970. 'Aboriginal settlements: a survey of 
institutional communities in eastern Australia'. Australian 
National University: Canberra. 

MADDOCK, K. 1972. The Australian Aborigines: a portrait 
of their society'. Allen Lane The Penguin Press: London. 

MCCARTHY, F. D. 1964. The dancers of Aurukun. 
Australian Natural History 14: 296-300. 

MCCARTHY, F. D. 1978. Aurukun dances. Typescript MS, 
3 parts, Sydney. Australian Institute of Aboriginal and 
Torres Strait Islander Studies Library: Canberra. 

McCONNEL, U. H. 1930. The Wik-munkan tribe of Cape 
York Peninsula, Part II. Oceania 1: 181-205. 

McCONNEL, U. H. 1953. Native arts and industries on the 
Archer, Kendall and Holroyd Rivers, Cape York 
Peninsula, North Queensland. Records of the South 
Australian Museum 11: 1—42. 

McCONNEL, U. H. 1957. 'Myths of the Mungkan'. 
Melbourne University Press: Melbourne. 

MORPHY, H. 1981. Art of northern Australia. Pp.53-65 in 
'Aboriginal Australia*. Ed. C. Cooper, H. Morphy, J. 
Mulvaney & N. Peterson. Australian Gallery Directors 
Council: Sydney. 

RHODES, E. G. 1980. Modes of holocene coastal 
progradation, Gulf of Carpentaria. PhD thesis, Australian 
National University. 

RIGSBY, B., & SUTTON, P. 1980-82. Speech communities 
in Aboriginal Australia. Anthropological Forum 5: 8-23. 



SHARP, R. L. 1940. An Australian Aboriginal population. 
Human Biology 12: 481-507. 

SUTTON, P. 1978. Wik: Aboriginal society, territory and 
language at Cape Keerweer, Cape York Peninsula, 
Australia. PhD thesis, University of Queensland. 

SUTTON, P. 1988. Dreamings. Pp.13-32. in 'Dreamings: 
The art of Aboriginal Australia 1 . Ed. P. Sutton. George 
Braziller: New York. 

SUTTON, P., MARTIN, D., von STURMER, J., CRIBB, R. 
& CHASE, A. 1990. 'Aak: Aboriginal estates and clans 
between the Embley and Edward Rivers, Cape York 
Peninsula'. South Australian Museum: Adelaide 
(restricted). 

SUTTON, P. & SMYTH, D. 1980. Ethnobotanical data from 
Aurukun (Queensland) area. Computer printout, 
Australian Institute of Aboriginal and Torres Strait 
Islander Studies Library, Canberra. 

THOMSON, D. F. 1932. Ceremonial presentation of fire in 
North Queensland. A preliminary note on the place of fire 
in primitive ritual. Man 32: 162-66. 

THOMSON, D. F. 1936. Notes on some bone and stone 
implements from North Queensland. Journal of the Royal 
Anthropological Institute 66: 71-74. 

THOMSON, D. F. 1939a. The seasonal factor in human 
culture illustrated from the life of a contemporary nomadic 
group. Proceedings of the Prehistoric Society 5: 209-21. 

THOMSON, D. F. 1939b. Notes on the smoking-pipes of 
North Queensland and the Northern Territory of Australia. 
Man 39: 81-91. 

von STURMER, J. R. 1978. The Wik region: Economy, 
territoriality and totemism in Western Cape York 
Peninsula, North Queensland. PhD thesis, University of 
Queensland. 



THE CHOOLKOONING METEORITE : A NEW (L6) OLIVINE 
HYPERSTHENE CHONDRITE FROM SOUTH AUSTRALIA 

M Zbik & A. Pring 



Summary 

The Choolkooning 001 meteorite is a single stone of 1.977 kg found approximately 63 km north of 
Hughes, South Australia, in 1991. It has been classified as an L6 chondrite shock facies S3-4 and 
contains olivine (Fa24.7), orthopyroxene (Fs20.5), plagioclase (Anl2 Or6.2Ab81.8), clinopyroxene, 
nickel-iron, troilite, chlorapatite and chromite. Mineral composition indicate that Choolkooning 001 
was a metamorphosed part of the L-planetoid and was moderately shocked before reaching Earth. 



THE CHOOLKOONING 001 METEORITE: A NEW (L6) OLIVINE-HYPERSTHENE 
CHONDRITE FROM SOUTH AUSTRALIA 



M.ZBIK&A.PRING 



ZBIK, M. & PRING, A. 1994. The Choolkooning 001 meteorite: a new (L6) olivine-hypersthene 
chondrite from South Australia. Rec. S. Aust. Mus. 27(1): 53-56. 

The Choolkooning 001 meteorite is a single stone of 1.977 kg found approximately 63 km north of 
Hughes, South Australia, in 1991. It has been classified as an L6 chondrite shock facies S3^ and 
contains olivine (Fa 247 ), orthopyroxene (Fs 2() ,.), plagioclase (An.-Gr ,Ab glg ), clinopyroxene, nickel- 
iron, troilite, chlorapatite and chromite. Mineral composition and textures indicate that Choolkooning 
001 was a metamorphosed part of the L-planetoid and was moderately shocked before reaching Earth. 

M. Zbik and A. Pring, Department of Mineralogy, South Australian Museum, North Terrace, Adelaide, 
South Australia 5000. Manuscript received 9 December, 1992. 



A single mass of the Choolkooning 001 meteorite, 
now in several pieces, was found by an unknown 
collector, 30 km west of Choolkooning Rockhole 
and 63 km north of Hughes on the South Australian 
part of the Nullarbor Plain. The approximate co- 
ordinates for the site are 29°55'S, 129°50'E. The 
broken and weathered fragments, total mass about 2 
kg, were illegally collected and exported to the 
United States of America in 1991. The meteorite 
was purchased in the United States of America by 
an Australian mineral dealer, Mr Robert Sielecki, 
who surrendered it to the South Australian Museum 
upon his return to Australia in March, 1992. 

In recent years the Nullarbor Plain has proved to 
be a productive area for the recovery of meteorites 
(Bevan 1992; Bevan & Binns 1989a, 1989b) and in 
the last few years has attracted the attention of 
illegal meteorite collectors. Under legislation 
enacted by the Governments of Western Australia 
and South Australia all meteorites found in these 
states are the property of the Crown. An unfortunate 
consequence of the illegal trade in meteorites is the 
loss of important information on the exact date and 
location of the find. The locality given for the 
Choolkooning 001 meteorite by the dealer from 
whom Mr Sielecki purchased the stone was 100 km 
north east of Deakin, Western Australia. This places 
the find site within South Australia, approximately 
63 km north of Hughes, some 30 km west of 
Choolkooning Rockhole (Fig. 1). In accordance with 
the guidelines on the nomenclature of meteorites 
from the South Australian Nullarbor (Bevan & Pring 
1993), the meteorite has been named Choolkooning 
001, being the first meteorite to be recorded from 
the Choolkooning area. 



Physical Description 

The meteorite is in eight fragments which range 
in size from 5 cm in length up to 20 cm. Four of the 
fragments fit together to form an incomplete 
rounded stone, two other fragments also appear to 
be part of this stone, but the other two fragments 
may be pieces from another mass of the meteorite. It 
is clear that several fragments are missing and it is 
not possible to say with certainty whether the 
meteorite was originally one or two stones. 

Each piece shows a dark brown 1 mm thick 
fusion crust on at least one surface. The stone is 
heavily weathered displaying iron staining of silicate 
minerals and fractures filled with iron oxides. The 
broken surfaces of the fragments are also heavily 
weathered and the fusion crusts of some fragments 
are encrusted with calcrete, indicating that the 
meteorite had been exposed to the weather for many 
years. The interior of the meteorite is grey in colour 
and medium to fine grained. 

In thin section the meteorite is generally light 
coloured with heavy iron oxide staining in some 
patches (Fig. 2). The chondrules are recrystallised 
and have poorly defined boundaries but are 
recognisable under crossed polars. They are 
typically 0.5 mm in diameter but some measuring 
up to 4 mm in diameter were also noted. Metal and 
troilite occurs as finely disseminated grains 
throughout the matrix. One piece of the meteorite 
was cut and the surface polished, and a polished 
thin section was also prepared and used for 
petrographic examination and in electron 
microprobe analyses. 



54 



M. ZBIK&A. PRING 



N 



Choolkooning 
Rockhole 



* 29 ° l Choolkooning 

Rockhole i 



130°' 




NULLARBOR 

i-T-r-T- rn-nr^ri irn 

PLAIN 




Gulf- 

1st 

Vincent 
Kangaroo Is. 



rv 1 




FIGURE 1. Map of South Australia showing the approximate location of the Choolkooning 001 meteorite. 



TABLE 1. Average chemical compositions of major minerals in the Choolkooning 001 meteorite. 



oxide 



olivine 



weight % 
ortho pyroxene 



plagioclase 



Si0 2 


39.1 


54.1 


63.8 


Ti0 2 


0.1 


0.2 


0.1 


A1 2 0, 


0.1 


0.1 


21.0 


FeO 


22.8 


14.7 


0.6 


MnO 


0.5 


0.5 


0.1 


MgO 


38.8 


28.8 


0.1 


CaO 


0.1 


0.7 


2.3 


Na 2 


- 


- 


8.5 


K.O 


- 


- 


1.0 


Cr 2 0, 


0.1 


0.1 


- 



NiO 



0.1 



Total 



101.7 



99.3 



97.5 



CHOOLKOONING METEORITE 



55 




FIGURE 2. Photomicrograph of the Choolkooning 001 meteorite in thin section showing typical fractured olivine grains in the 
re crystallized matrix. 



Mineralogy 

Compositions of the silicate minerals were 
determined with a JEOL electron microprobe at the 
University of Adelaide Centre for Electron 
Microscopy and Microbeam Analysis. Analyses 
were made using an accelerating voltage of 15 kV, a 
sample current of 3 nA, and a beam width of 5 urn. 
Representative mineral analyses are presented in 
Table 1. 

Relic chondrules and chondrule fragments are 
composed predominantly of olivine, with lesser 
amounts of orthopyroxene. 'Barred' chondrules 
composed of olivine and orthopyroxene also have 
thin lamellae of either clinopyroxene or plagioclase. 
The matrix consists of olivine and orthopyroxene 
grains with very rare grains of clinopyroxene up to 
20 urn in diameter. Plagioclase is abundant 
throughout the matrix as small turbid grains which 
display undulate extinction. Coarser (up to 500 um) 
grains of plagioclase feldspar also occur in the 
matrix with some exhibiting albite twinning. 
Nickel-iron metal, troilite, chlorapatite and chromite 
occur as accessory minerals. 

Microprobe analyses show that the olivine in the 
Choolkooning 001 meteorite is equilibrated with a 



mean fayalite content of Fa 247 (range Fa 23g 254 , 30 
analyses). The orthopyroxene shows only a small 
variation in chemical composition with a mean 
ferrosilite content of Fs 203 ( 30 analyses) and a 
wollastonite content of 1.5 mol%. Clinopyroxene 
was identified but the grains were too small to 
provide reliable analyses. The plagioclase content 
was found to be An 12 Or 62 Ab g| g (1 1 analyses). 

The pyroxene geothermometers of Wells (1977) 
and Lindsley (1983) suggest that the Choolkooning 
001 meteorite was heated to temperatures of 
between 700° and 800° C during metamorphism 
while the meteorite was still part of a large L type 
asteroid. The Choolkooning 001 meteorite is very 
similar in composition to the Mangalo meteorite, an 
L6, recently described from Eyre Peninsula 
(Wallace & Pring 1991). 



Classification 

The Choolkooning 001 meteorite has been 
classified as an L6 chondrite. The olivine 
composition is within the range of the L chondrites 
(Keil & Fredriksson 1964), and the orthopyroxene 
composition (Fs 205 ) shows that the meteorite 



56 



M. ZBIK & A. PRING 



belongs to the olivine-hypersthene chondrite group. 
The highly equilibrated chemical composition, 
crystalline matrix, poorly defined chondrule 
boundaries, well recrystallized plagioclase and 
euhedral chlorapatite crystals suggest that 
Choolkooning 001 belongs to the type 6 
classification of Van Schmus & Wood (1967). The 
wollastonite content in the orthopyroxene is similar 
to that found in most L6 chondrites (Scott et al. 
1986). The presence and content of plagioclase, the 
degree of crystal fracture and the undulate and 
mosaic extinction with planar fractures all indicate 
that the meteorite has been moderately shocked after 
metamorphism. According to the classification 



scheme of Stoffler et al. (1991) the shock facies is 
estimated to be S3^t, moderately shocked. 



Acknowledgments 

The authors wish to thank Mr G. Horr for preparing a 
polished thin section of the meteorite and Mr Huw Rosser of 
CEMMA, University of Adelaide for assistance with the 
electron microprobe analyses. The financial support of the Ian 
Potter Foundation and the Friends of the South Australian 
Museum is gratefully acknowledged. We also wish to thank 
Dr. Alex Bevan and Dr. Margaret Wallace for their 
constructive comments on the manuscript. 



References 



BEVAN, A. W. R. 1992. Australian Meteorites. Records of 
the Australian Museum, Supplement 15: 1-27. 

BEVAN , A. W. R. & BINNS, R. A. 1989a. Meteorites from 
the Nullarbor Region, Western Australia: I. A review of 
past recoveries and a procedure for naming new finds. 
Meteoritics 24: 127-133. 

BEVAN, A. W. R. & BINNS, R. A. 1989b. Meteorites from 
the Nullarbor Region, Western Australia: II. Recovery and 
classification of 34 new meteorite finds from the 
Mundrabilla, Forrest, Reid and Deakin areas. Meteoritics 
24: 135-141. 

BEVAN, A. & PRING, A. 1993. Guidelines for the 
nomenclature of meteorites from the South Australian 
Nullarbor. Meteoritics 28: 600-602. 

KEIL, K. & FREDRIKSSON, K. 1964. The iron, magnesium 
and calcium distributions in coexisting olivines and 
rhombic pyroxenes of chondrites Journal of Geophysical 
Research 69: 3487-3515. 

LINDSLEY, D. H. 1983. Pyroxene thermometry. American 
Mineralogist 68: 477-493. 



SCOTT, E. R. D„ TAYLOR, G. J. & KEIL, K. 1986. 
Accretion, metamorphism, and brecciation of ordinary 
chondrites: evidence from petrologic studies of meteorites 
from Roosevelt County, New Mexico. Proceeding of the 
Lunar and Planetary Science Conference 17: El 15— 
E123. 

STOFFLER, D., KEIL, K. & SCOTT, E. R. D. 1991. Shock 
metamorphism of odrdinary chondrites. Geochimica 
Cosmochimica Acta 55: 3845-3867. 

VAN SCHMUS, W. R. & WOOD, J. A. 1967. A chemical- 
petrologic classification for the chondritic meteorites. 
Geochimica Cosmochimica Acta 31: 747-765. 

WALLACE, M. E. & PRING, A. 1991. The Mangalo 
meteorite, a new (L6) olivine-hypersthene chondrite from 
South Australia. Transactions of the Royal Society of 
South Australia 115: 89-91. 

WELLS, P. R. A. 1977. Pyroxene thermometry in simple and 
complex systems. Contributions to Mineralogy and 
Petrology 62: 129-139. 



OBITUARY : SHANE ALWYN PARKER 3 AUGUST 1943 - 21 NOVEMBER 

1992 

W. Zeideler 



Summary 

This obituary gives emphasis to Shane's work in marine invertebrates and lists all of his 
publicatons. His ornothological work was emphasised by Horton (1993). 



OBITUARY 



SHANE ALWYNE PARKER 



3 August 1943 - 21 November 1992 




This obituary gives emphasis to Shane's work in 
marine invertebrates and lists all his publications. 
His ornithological work was emphasised by Horton 
(1993). 

Shane Alwyne Parker was born in Colchester, 
Essex where he attended schools gaining his O- 
Level in seven subjects. His interest in natural 
history developed early with walks in the English 
countryside with his aunt. He was an obsessive 
collector with an inquiring mind and often visited 
the Colchester Museum to identify natural history 
specimens that he had collected. Sadly, Shane's 
father did not approve of this "obscure" interest and, 
as Shane related to me, his life at home was not a 
happy one. At the age of 16 he began work at the 
British Museum, at Bloomsbury, initially as 
assistant in the Photographic Section, Reading and 
Map Room and later as assistant in the Bird Section 
of the Natural History Museum. He used to travel 
on the train from Colchester to London and back 
every day, leaving home at some ungodly hour each 
morning. Sometime later he moved to London and 
lived in a hostel where he obviously had a very 
merry time. He spent six and a half years at the 
British Museum, three as curator of their egg 
collection. In 1964 he took part in the second of the 
Harold Hall (British Museum) expeditions in 
Australia to northern Queensland from February to 
August. Those seven months were a highlight in 
Shane's life and influenced him to emigrate to 
Australia in 1967. 



From 1967 to 1971 Shane worked for the Arid 
Zone Research Institute, Alice Springs, first as 
assistant to the Biologist and then to the Botanist, 
taking part in faunal and floral surveys of the 
Northern Territory and curating zoological and 
botanical collections. During this period (January - 
March, 1968) Shane undertook a solo trip to 
Choiseul and Malaita, Solomon Islands, in search of 
a species of ground-pigeon, without success (40). 
Later, working on a wallaby survey (39), he moved 
to Darwin where he met his future wife, Erica, 
whom he married in 1970 in Alice Springs. They 
moved to Adelaide in 1971 where Shane, feeling 
the need for a formal education, studied full-time at 
Daws Road High School to gain his Matriculation in 
1972. He went on to complete a Bachelor of Science 
degree, majoring in Botany and Zoology, at the 
University of Adelaide in 1975. 

During all this time Shane's lack of formal 
qualifications did not prevent him from publishing 
his observations. He published his first paper at the 
age of 16 (1) and by the end of 1967, while at the 
British Museum, had published 24 notes and 
articles, probably embarrassing some of his 
colleagues who encouraged him to emigrate. During 
his early years in Australia he continued to publish 
ornithological observations, corresponding regularly 
with other workers in Australia and overseas and 
while in Adelaide became an Honorary Research 
Worker at the South Australian Museum. When 
Herbert Condon retired as Curator of Birds in late 



58 



W ZEIDLER 



1974 Shane, while completing his final year at 
University, was asked to run the Bird Section until 
a replacement could be found. By this time Shane 
had published a further 25 articles including one on 
plants (38) and two on mammals (39, 47), the latter 
publication being a significant contribution. 
Logically and fittingly Shane was appointed Curator 
of Birds in January 1976 - a position which he held 
full-time until 1985 when he began his work as 
Curator of Lower Marine Invertebrates. This dual 
role continued until October 1991 although his work 
in ornithology consisted mainly of completing small 
research projects and training Dr Philippa Horton to 
take over the management of the Bird Section, the 
position of 'Curator of Birds' having been 
abolished. Tragically, by this time, Shane had been 
diagnosed with a lymphoma and although initial 
treatment seemed successful he lost his fight of 
about two years and died peacefully at home on 21 
November 1992. 

During his relatively brief lifetime Shane 
published some 118 notes and articles (including 
seven book reviews), more than 100 of them on 
ornithology. There is no doubt that Shane's 
knowledge of birds of the world and the 
Australasian bird fauna in particular was highly 
regarded worldwide, and it was for this reason that 
he was chosen as one of only five Australian 
delegates to visit China in 1981 under the auspices 
of the Australian and Chinese Museums Association 
(83). He was an active member of the South 
Australian Ornithological Association serving as 
Vice President in 1979-81 and 1985-86. He also 
served the Royal Australian Ornithologists Union as 
Councillor (1978-81), Chairman, Record Appraisal 
Committee (1979-82) and on the Taxonomic 
Advisory Committee (1979-83); and in 1978-81 he 
was on the Conservation Programme Committee of 
the World Wildlife Fund (Australia). He was also 
Secretary for the Royal Society of South Australia 
for the year 1986-87. 

Amongst ornithologists Shane will best be 
remembered for his contribution to the Museum's 
bird collections. With the help of a dedicated band 
of volunteers, which he cultivated, Shane began the 
process of modernising the collection. Almost from 
scratch he began to establish parallel collections of 
skeletons, whole spirit specimens, nests and 
stomach contents. He conducted field work mainly 
to collect downy young and immatures of non- 
passerines, domestic species and neglected common 
species, and further augmented the collection and 
filled in gaps by exchanges with other museums. As 
a result of those activities the bird collections 
increased dramatically in size and scientific 
significance. In a relatively short time Shane, with 
his acquisition program and meticulous 
documentation, had transformed the Museum's bird 



collections from one used mainly by bird watchers 
to one of science. He was always generous with his 
time making himself and the collection very 
available to bird enthusiasts and spent many hours 
encouraging young people to develop their interest 
in and knowledge of birds. Many of these people 
are now employed in the field of natural science and 
environmental studies. A highlight of his 
curatorship was the good relationship which he 
cultivated with the descendants of Captain S. A. 
White resulting in the donation to the Museum of 
White's African bird skin collections in 1976 and 
the main skin and egg collection in 1988. Shane's 
relationship with the White family culminated in 
the Museum's exhibition 'Captain White and the 
Hcuse of Birds'. Regrettably he died only 24 days 
before the exhibition opened. 

Shane delighted in detective work, whether it was 
unravelling details of historical collecting 
expeditions and collections (18, 71, 91, 92, 108) or 
taxonomic problems (42, 88, 90). He also loved 
searching for 'lost' species (40, 62) and in 1979 
joined an expedition on camel to search for the night 
parrot around Cooper's Creek. He believed as the 
only known specimens were discovered on camel he 
was more likely to succeed by these means if the 
bird was still extant. The party sighted four birds 
east of Lake Perigundi but Shane was not entirely 
convinced and never published a detailed account of 
the observations. 

For his ornithological work Shane was honoured 
by the naming of a new subspecies of Honeyeater, 
Acanthogenys rufogularis parkeri (Parkes 1980) 
and two species of feather louse from the mallee 
fowl (Price & Emerson 1984; Emerson & Price 
1986). On the latter Shane remarked "to the delight 
of my friends and the even greater delight of my 
enemies!". Although his career in Marine 
Invertebrates was brief he touched many with his 
enthusiasm, clarity of thought and meticulous 
research and his memory will be honoured by the 
naming of several species by various colleagues in 
future publications. 

During the early 1980s Shane had some 
unfortunate disagreements with a few amateur 
ornithologists and his over-sensitive nature led him 
into deeper conflicts than need have been. These 
arguments came to a head over the description of a 
new sandpiper (79) resulting in a prolonged conflict 
and threatened legal action by both parties. These 
events were particularly distressing for Shane as he 
prided himself on his honesty and integrity and it 
hurt him deeply when he did not receive the support 
from his colleagues that he might have expected. In 
addition Shane was facing ill-informed criticism 
from a few amateur ornithologists and the public for 
the collection of specimens, particularly in regard to 
the Eyrean grass wren. Shane ultimately became very 



OBITUARY - SHANE A. PARKER 



59 



disillusioned with the Australian ornithological 
community and in 1985 he switched his energies to 
the much neglected lower marine invertebrates. 

Observing Shane working in the Marine 
Invertebrates Section was like watching a child with 
new found toys. There were just so many areas 
where he could apply his love for classical 
taxonomy. To his surprise and delight he found that 
new species could be found almost on a daily basis 
and new genera and families lurked in the collection 
or amongst newly acquired material from deep-sea 
trawlers. 

Confronted with an array of animal groups, some 
of which he didn't even realise existed, he quickly 
sought out experts and corresponded with them. 
Much to his surprise he found them only too willing 
to help and share information. He corresponded 
often and developed a relationship with some 
colleagues that resulted in several joint publications 
describing the fauna of southern Australia. One of 
his major articles, and one of which he was 
justifiably proud, was a monograph on the stony 
corals of South Australia, Victoria and Tasmania 
which he produced with S. D. Cairns of the 
Smithsonian Institution as senior author (111). It 
was his last paper to be published before his death. 
Shane also published on polychaetes and bryozoans 
and at the time of his death was working on a couple 
of papers on leeches. 

I was continually amazed by Shane's ability to 
comprehend the complex systematics of some phyla, 
if only to come up with a scheme of classification so 
that he could properly curate the collections. He 
very quickly identified specimens as best he could, 
completely reorganised the collections and curated 
them according to the most modern, acceptable 
classification. Whenever he came across a group for 
which he could not find an acceptable scheme of 
classification, he would write to one of the experts, 
usually overseas, and ask for help. Often this kind 
of correspondence led to the material being loaned 
for study, resulting in publications describing the 
unique fauna of southern Australia. Shane 
encouraged the use of the collections by others and 
was very generous with his time, often humbly 
accepting junior authorship for his efforts. He 
organised many exchanges of specimens and added 
substantially to the literature base of the collections. 
Shane was rewarded for his curatorial efforts when 
an overseas colleague, in a submission to the 
Museum Review 1991/92, remarked 'I have worked 
extensively at the U.S. National Museum, 
Smithsonian Institution and also at the British 
Museum and the collections at the South Australian 
Museum are far superior in quality'. These remarks 
and ones in a similar vein from other colleagues 
pleased Shane very much. 

Shane soon came to realise that he could not do 



everything and I encouraged him to specialise in 
polychaetes or bryozoans. It was the latter group 
that eventually 'grabbed' him and he was 
continually amazed by the diversity and beauty of 
the little 'critters'. Shane was never anyone to do 
things by halves and even when he was ill he was 
working on his checklist of Australian bryozoans 
which was to form the basis for future major 
taxonomic revisions of the entire Australian fauna. 
Fortunately for bryozoan workers Shane's checklist 
will be completed by his colleague and collaborator, 
Dr Peter Haywood, University of Wales, Swansea. 
But sadly, a wealth of knowledge passed away with 
Shane and we will never know the greatness that he 
was sure to achieve. 

To some people, particularly to those who did not 
know him well, Shane was an English eccentric 
born about a century too late. He was a great 
admirer of Conan Doyle's hero, Sherlock Holmes 
and in winter often wore a tailor-made Edwardian 
frock-coat and wing collars. Although Shane loved 
elegant and stylish clothes, particularly the 
Edwardian fashions, and may have admired the life 
style of Edwardian society, that did not necessarily 
make him old fashioned. He was quite up with 
modern technology and methods and was 
instrumental in the purchase of a Scanning Electron 
Microscope for the Museum and used the 
instrument regularly for his research. However, I 
think he may have been more comfortable with 
computers had they been steam driven. Shane was 
widely read especially in ancient history, natural 
history and exploration and had a broad and deep 
knowledge of a wide variety of subjects. He was 
skilled at languages, took lessons in Gaelic and 
taught himself Latin while commuting on the train 
so that he could read the classics in the original. He 
also enjoyed a good detective story and was partial 
to the obscure and bizarre with Dan Dare comics, 
Winnie the Pooh, Molesworth and 'William* books 
being some of his favourites. He was an 
acknowledged authority on bawdy English folksongs 
which he could sing well in the appropriate accent. 
His collection of songs is regarded by some as one 
of the best in the world, valued for its academic 
content as well as for the many rare and original 
versions. Shane also loved fine music whether it 
was classical or modern with Pachelbefs Canon in 
D being one of his favourites. He was a talented 
artist and purchased his first bird book with money 
from selling his bird paintings to relatives and 
friends. He was particularly good at caricature and 
comic strips which provided an outlet for his 
wonderful, unique sense of humour. He enjoyed 
working in his vegetable garden and amongst his 
many other interests was that of steam trains and 
collecting finely engraved stamps and cigarette 
cards. 



60 



W. ZEIDLER 



Perhaps his greatest passion was that of good food 
and wine. He was sometimes 'caught' in his office 
devouring some great custard pie or cream bun. Yet 
these gastronomic excesses had little effect on his 
slight frame. In the Marine Invertebrates Section, 
cream buns or preferably, huge creamy cakes, were 
compulsory on birthdays and whenever a paper was 
published - we seemed to have rather a lot of those! 
Shane was a founding member and the driving force 
behind the Museum select dining club De Gustibus 
and was the editor of its occasional organ The 
Gullet. This gave him a wonderful opportunity to 
make use of his creative writing skills. He loved the 
English language and took great delight in using it. 
He spent hours over the articles and the presentation 
of The Gullet and kept copies of all the issues. 
Perhaps as a result of his trip to China in 1981, 
where he attended many banquets of duck's feet and 
other unmentionable or unidentifiable gastronomic 
delights, it was the charter of De Gustibus to dine 
out at unusual places and sample unusual dishes. 
The annual De Gustibus Christmas Luncheon was a 
grand affair. It was the duty of everyone attending to 
provide an exotic dish that they had prepared 
together with good wine and champagne. Shane 
would address the meeting apologising for the 
absence of our fictitious president Sir Gregory 
Parsloe-Parsloe K.B. (Shane's alter-ego) who was 
unable to attend because of some hilarious and often 
disgusting, gluttonous mishap. The whole 
proceedings were always accompanied by live 
chamber music with Pachelbel's Canon being 
played at the beginning and end. For me these 
annual events were some of the most memorable of 
my life and I am grateful to Shane for the 
experience. Members of De Gustibus will be 
pleased to learn that before his death Shane 
honoured Sir Gregory by the naming of an 
appropriately large bryozoan; let us hope it gets 
published before the editor finds out! I will miss his 
good humour and sharp wit and I will never forget 
his last words to me as he said with a mischievous 
smile and knowing glint in his eye 'See you anon 
old bean'. 

To Shane's wife Erica and their sons Gathorne 
and Tolle we extend our heartfelt sympathy. 

Bibliography 

19 60 

1. Snail-eating in the redbacked shrike (Lanius 
collurio). Essex Naturalist 30: 276-278. 



1962 

2. A new name for Bradypterus barratti major 
(Roberts). Bulletin of the British Ornithologists' 
Club 82: 122. 

3. Notes on some undescribed eggs from New Guinea. 



Bulletin of the British Ornithologists' Club 82: 132- 
133. 



1963 



4. The validity of the genus Lusciniola Gray. Bulletin 
of the British Ornithologists' Club 83: 65-69. (With 
C. J. O. Harrison as junior author). 

5. Nidification of the genus Melanocharis Sclater 
(Dicaeidae). Bulletin of the British Ornithologists' 
Club S3: 109-112. 

6. Nesting of the paradise crow, Lycocorax 
pyrrhopterus (Bonaparte) and the spangled drongo, 
Dicrurus hottentottus (Linn.) in the Moluccas. 
Bulletin of the British Ornithologists' Club 83: J 26— 
127. 

7. A note on the habits of Mayrornis schistaceus Mayr 
(Muscicapidae) of the South-West Pacific. Bulletin of 
the British Ornithologists' Club 83: 159-161. 

8. Eggs of Francolinus africanus lorti and 
Creatophora cinerea from Somaliland. Oologist's 
Record 37: 41-42. 

1964 

9. Taxonomic position of the genus Culicicapa Swinhoe 
(Muscicapidae). Bulletin of the British 
Ornithologists' Club 84: 45-46. 

10. The identity of Antiornis grahami Riley. Bulletin of 
the British Ornithologists' Club 84: 113-1 14. 

1965 

11. The eggs of the white-throated greenbul 
Phyllastrephus albigularis (Sharpe). Bulletin of the 
British Ornithologists' Club 85: 95. (With C. J. O. 
Harrison as senior author). 



12. 



13 



Some misidentified eggs of the grey wood-shrike 
Tephrodornis gularis (Raffles) and the long-tailed 
sibia Heterophasia picaoides (Hodgson). Bulletin of 
the British Ornithologists' Club 85: 96. (With C. J. 
O. Harrison as senior author). 



Fieldnotes on the birds of the Santa Cruz Islands, 
south-west Pacific. Bulletin of the British 
Ornithologists' Club 85: 154-159. (With C. J. 
Hadley as senior author). 

14. The behavioural affinities of the blue wrens of the 
genus Malurus. Emu 65: 103-113. (With C. J. O. 
Harrison as senior author). 

1966 

15. The taxonomic affinities of the New Guinea genera 
Paramythia and Oreocharis, Bulletin of the British 
Ornithologists' Club 86: 15-20. (With C. J. O. 
Harrison as senior author). 

16. The eggs of the white-tailed blue chat, Cinclidium 
leucurum, and the Large Niltava, Niltava grandis. 
Bulletin of the British Ornithologists' Club 86: 71- 
73. (With C. J. O. Harrison as senior author). 

1 7. Coracias abyssinica: an unrecorded host of Indicator 
indicator. Bulletin of the British Ornithologists' 
ClubS6: 81. 



OBITUARY - SHANE A. PARKER 



61 



18. Albert S. Meek's collecting-locality on the Cape 
Yorke Peninsula, 1898. Emu 66: 121-122. 

1967 

19. The eggs of Woodford's rail, Rouget's rail, and the 
Malayan banded crake. Bulletin of the British 
Ornithologists' Club 87: 14-16. (With C. J. O. 
Harrison as senior author). 

20. New information on the Solomon Islands crowned 
pigeon, Microgoura meeki Rothschild. Bulletin of the 
British Ornithologists' Club 87: 86-89. 

21. Some eggs from the New Hebrides, south-west 
Pacific. Bulletin of the British Ornithologists' Club 
87:90-91. 

22. The eggs of the wattled brush turkey Aepypodius 
arfakianus (Salvadori) (Megapodiidae). Bulletin of 
the British Ornithologists' Club 87: 92. 

23. AS. Meek's three expeditions to the Solomon Islands. 
Bulletin of the British Ornithologists' Club 87: 129- 
135. 



24. 



1968 

25. 



26 



The identification of the eggs of Indian Hill Partridges 
of the genus Arborophila. Journal of the Bombay 
Natural History Society 63: 748-750." (With C. J. O. 
Harrison as senior author). 



An instance of apparent sympatry between the great 
and spotted bowerbirds. Bulletin of the British 
Ornithologists' Club 88: 56. 



An instance of apparent sympatry between the great 
bowerbird and the spotted bowerbird. Emu 68: 222. 

27. The type-locality of the White-quilled Rock Pigeon, 
Petrophassa albipennis Gould. Bulletin of the 
British Ornithologists' Club 88: 57-58. 

28. On the thick-billed ground dove Gallicolumba 
salamonis (Ramsay). Bulletin of the British 
Ornithologists' Club 88: 58-59. 

1969 

29. New and interesting distribution records of Central 
Australian birds. South Australian Ornithologist 25: 
59-71. 

30. The Atherton scrub-wren Sericornis keri Mathews, a 
neglected Australian species. Emu 69: 212-232. 
(With I. C. J. Galbraith as senior author). 

1970 

3 1 . Critical notes on the status of some Northern Territory 
birds. South Australian Ornithologist 25: 1 15-125. 

32. Taxonomy of the populations of Sericornis beccarii 
inhabiting Cape Yorke Peninsula. Emu 70: 69-72. 

1971 

33. Taxonomy of the Northern Territory friarbirds known 
as Philemon buceroides gordoni. Emu 71: 54-56. 

34. Association betwen the sulphur-crested cockatoo and 
Pandanus. Western Australian Naturalist 12: 23. 



35. Critical notes on the status of some central Australian 
birds. Emu 71: 99-102. 

36. First record of the yellow wagtail in the Northern 
Territory. Emu 71: 142. (With D. N. Crawford as 
senior author). 

37 . Distribution of Meliphaga flavescens and M. fusca in 
northern Queensland. Sunbird 2: 41-47. 

38. Further collections of two little-known Stylidiaceae 
from the Northern Territory. Muelleria 2(2): 145- 
146. (With J. R. Maconochie as senior author). 

39. Notes on the small black wallaroo Macropus 
bernardus (Rothschild, 1904) of Arnhem Land. 
Victorian Naturalist 38: 41-43. 

1972 

40. An unsuccessful search for the Solomon Islands 
crowned pigeon. Emu 72: 24-26. 

41. The occurrence of the yellow chat in south-western 
Queensland. Sunbird 3: 15. (With J. R. Ford as senior 
author). 

42. Remarks on distribution and taxonomy of the grass 
wrens Amytornis textilis, modestus and purnelli. 
Emu 72: 157-166. 

1973 

43. The tongues of Ephthianura and Ashbyia. Emu 73: 
19-20. 

44. The identity of Microeca brunneicauda Campbell, 
1902. Emu 73: 23-25. 

45. First record of Acanthiza robustirostris in 
Queensland. Emu 73: 27. (With J. R. Ford as senior 
author). 

46. A second species of wedgebill? Emu 73: 113-118. 
(With J. R. Ford as senior author). 

47. An annotated checklist of the native land mammals of 
the Northern Territory. Records of the South 
Australian Museum 16(1 1): 1-57. 

1974 

48. Distribution and taxonomy of some birds from south- 
western Queensland. Emu 74: 177-194. (With J. R. 
Ford as senior author). 

1975 

49. Maluridae. P. 1 1 in 'Interim List of Australian 
Songbirds'. R. Schodde. Royal Australasian 
Ornithologists' Union: Melbourne. 

1976 

50. Some notes on the birds of Kangaroo Island. South 
Australian Ornithologist 27: 123-124. (With A. F. 
C. Lashmar as junior author). 

51. The occurrence of the singing bushlark Mirafra 
javanica in north-eastern South Australia. South 
Australian Ornithologist 27: 143. (With P. F. 
Lawson as senior author). 

52. Further notes on the birds of Groote Eylandt, N.T. 



62 



W. ZEIDLER 



South Australian Ornithologist 27: 144. (With G. 
M. Storr as junior author). 

53. First record of the erect-crested penguin Eudyptes 
sclateri from South Australia. South Australian 
Ornithologist 27: 146-147. (With B. R. Hutchins as 
senior author). 

54. Stipiturus, Amytornis, Ephthianura, Ashbyia. Pp. 
416-421, 510-513 in 'Reader's Digest Complete 
Book of Australian Birds'. Reader's Digest Services 
Pry Ltd: Sydney. 

1977 

55. Notes on the distribution of the chiming and 
chirruping wedgebills in South Australia. South 
Australian Ornithologist 21'. 175-176. 

56. Records of the barking owl from South Australia. 

South Australian Ornithologist 27: 204-206. 

57. The distribution and occurrence in South Australia of 
owls of the genus Tyto. South Australian 
Ornithologist 27: 207-215. 

58. Birds as pollinators. Australian Bird Watcher 7: 
124-133. 

59. Comment on the collecting of birds for taxonomic 
research. South Australian Ornithologist Association 
Newsletter September 83: 8-11. 

60. Further comments on collecting. South Australian 
Ornithologist Association Newsletter, December 84: 
4-8. 

1978 

61. Recommended English names for Australian birds. 
Emu 11 (supplement): 245-307. (With R. Schodde 
as senior author, and four other co-authors). 

62. Some observations on the Eyrean grasswren 
Amytornis goyderi (Gould, 1875). Records of the 
South Australian Museum 17(24): 361-371. (With 
I. A. May and W. Head as junior authors). 

63. Comments on some criticisms of the Interim List of 
Australian Songbirds. Australian Birds 13: 70-7 1 . 

64. Notes on the birds of Pearson, Dorothee and Greenly 
Islands, South Australia. Transactions of the Royal 
Society of South Australia 102: 191-202. (With J. 

B. Cox as junior author). 

65. Pedionomidae, Climacteridae, Dicaeidae, 
Ephthianuridae. Pp. 81-82, 222-223, 224-226, 230 
in 'Bird Families of the World'. Ed. C. J. O. Harrison. 
Elsevier-Phaidon: Oxford. 

1979 

66. 'An Annotated Checklist of the Birds of South 
Australia. Part One: Emus to Spoonbills'. South 
Australian Ornithological Association: Adelaide. 
(With H. J. Eckert, G. B. Ragless, J. B. Cox and N. 

C. H. Reid as junior authors). 

67. Remarks on the status of some Australian passerines 
Pp. 109-115 in 'The Status of Endangered 
Australasian Wildlife'. Ed. M. J. Tyler. Royal Society 
of South Australia: Adelaide. (With N. C. H. Reid as 
junior author). 



1980 

68. The records of the speckled warbler from South 
Australia. South Australian Ornithologist 28: 102- 
103. 

69. Birds and conservation parks in the north-east of 
South Australia. South Australian Parks and 
Conservation 3: 11-18. 

70. The spotted bowerbird in southwestern Queensland. 
Sunbird 10: 67-68. 

71. Samuel White's ornithological explorations in 
northern South Australia in 1863. South Australian 
Ornithologist 28: 113-119. 

72. An early breeding record of the sooty tern from 
Kangaroo Island. South Australian Ornithologist 28: 
139-140. 



1981 

73. 



74 



The status of the elegant parrot and the rock parrot on 
Kangaroo Island. South Australian Ornithologist 28: 
164-165. (With C. Baxter as senior author). 



Possible nest- parasitism in the Australian stiff-tailed 
ducks (Anatidae: Oxyurini). Emu 81: 41-42. (With 
A. R. Attiwill as senior author and J. M. Bourne as 
junior co-author). 

75. Prolegomenon to further studies in the Chrysococcyx 
'malayanus' group (Aves, Cuculidae). Zoologische 
Verhandelingen No. 187: 1-56. 

1982 

76. The relationships of the A ustralo- Papuan treecreepers 
and sittellas. South Australian Ornithologist 28: 
197-200. 

77. The records of the glossy black cockatoo from the 
South-East of South Australia. South Australian 
Ornithologist 28: 209-210. 

78. Additional notes on seabirds recorded in South 
Australia. South Australian Ornithologist 28: 213- 
216. (With I. A. May as junior author). 

79. A new sandpiper of the genus Calidris. South 
Australian Naturalist 56: 63. 

80. Notes on Amytornis striatus merrotsyi Mellor, a 
subspecies of the striated grasswren inhabiting the 
Flinders Ranges. South Australian Ornithologist 29: 
13-16. 

81. Remarks on the tympanic cavity of Malurus, 
Stipiturus and Amytornis (Passeriformes, Maluridae). 
South Australian Ornithologist 29: 17-22. 

82. The Bird section of the South Australian Museum - 
its collections and functions. Bird Care and 
Conservation Society Newsletter 12. 

83. Museums in China: a report by a delegation from the 
Museums Association of Australia. Pp. 63. (With P. 
Stanbury (Ed.), B. Bertram, D. Kitchener and B. 

Reynolds). 



1983 

84. 



Remarks on the taxonomy of the genus Calamanthus 
(fieldwrens). South Australian Ornithologist 29: 65- 
71. (With H. J. Eckert as junior author). 



OBITUARY - SHANE A. PARKER 



63 



85. Birds. Pp. 135-150 in 'Natural History of the South 
East'. Ed. M.J. Tyler et al. Royal Society of South 
Australia: Adelaide. (With N. C. H. Reid as junior 
author). 

86. Birds. Pp. 33-36 in 'Flinders Ranges National Park 
Management Plan'. Department of Environment and 
Planning: Adelaide. 

1984 

87. The relationships of the Madagascan genus 
Dromaeocercus (Sylviidae). Bulletin of the British 
Ornithologists' Club 104: 11-18. 

88. The extinct Kangaroo Island emu, a hitherto- 
unrecognized species. Bulletin of the British 
Ornithologists' Club 104: 19-22. 

89. Remarks on Struthidea cinerea dalyi Mathews. 
Sunbird 14: 7-lQ. 

90. The identity of Sericornis tyrannula De Vis. Emu 
84: 108-110. 

91. The African bird collections of S.A. White of South 
Australia. Scopus 8: 33-36. (With R. K. Brooke as 
senior author). 

92. Remarks on some results of John Gould's visit to 
South Australia in 1839. South Australian 
Ornithologist 29: 109-112. 

1985 

93. The occurrence of Lewin's rail on Eyre Peninsula, 
South Australia. South Australian Ornithologist 29: 
188. 

94. 'An Annotated Checklist of the Birds of South 
Australia. Part 2A: Waterfowl'. South Australian 
Ornithological Association: Adelaide. (With H. J. 
Eckert and G. B. Ragless as junior authors). 

95. Birds. Pp. 20-55 in 'A List of the Vertebrates of 
South Australia'. Edn 1. Ed. H. J. Aslin. Biological 
Survey Coordinating Committee and Department of 
Environment and Planning: Adelaide. 

96. First record of the slender-billed thornbill from the 
South-East of South Australia. South Australian 
Ornithologist 29: 212. 

97. Birds. Pp. 149-157 in 'Natural History of Eyre 
Peninsula'. Ed. C. R. Twidale et al. Royal Society of 
South Australia: Adelaide. (With H. J. Eckert as 
senior author and J. R. W. Reid as junior co-author). 

98. The rediscovery and taxonomic relationships of 
Gerygone igata amalia Meise, 1931. Bulletin of the 
British Ornithologists' Club 105: 118-121. 

99. Ephthianuridae, Rhipiduridae, Monarchidae, 
Acanthizidae, Maluridae. Pp. 85-86, 205-206, 358- 
359, 643-644, 664-665 in 'A Dictionary of Birds'. 
Ed. B. Campbell & E. Lack. T. and A. D. Poyser Ltd: 
Calton. 

1986 

100. First breeding record of the fleshy-footed shearwater 
in South Australia. South Australian Ornithologist 
30: 13-14. (With A. C. Robinson as senior author 
and A. Spiers as junior co-author). 



101 



1988 



A second specimen of the rufous fantail from South 
Australia. South Australian Ornithologist 30: 23. 
(With H. Bakker as senior author). 



102. The origin of the populations of the eastern rosella 
inhabiting the Mount Lofty Ranges and Adelaide 
Plains, South Australia. South Australian 
Ornithologist 30: 132. 



1990 

103. Birds. Pp. 21-54 in 'A List of the Vertebrates of 
South Australia'. Edn 2. Ed. C. H. S. Watts. 
Biological Survey Coordinating Committee and 
Department of Environment and Planning: Adelaide. 
(With P. Horton as junior author). 

104. First Australian records of the family Pisionidae 
(Polychaeta), with the description of a new species. 
Transactions of the Royal Society of South Australia 
114(4): 195-201. (With G. Hartmann-Schrbder as 
senior author). 



105. 



1991 



109. 



First Australian records of Hesionura (Polychaeta: 
Phyllodocidae), with the description of a new species. 
Transactions of the Royal Society of South Australia 
114(4): 203-205. (With G. Hartmann- Schroder as 
senior author). 



106. The plectriform apparatus - an enigmatic structure in 
malacostegine Bryozoa. Pp. 133-145 in Bigey, F. P. 
& d'Hondt, J.-L. (Eds) 'Bryozoa Living and Fossil'. 
Bulletin de la Societe des Sciences Naturelles de 
VOuest de la France (Nantes), Memoire HS 1: xiv + 
599 pp. (With D. P. Gordon as senior author). 

107. A new genus of the bryozoan family Electridae, with 
a plectriform apparatus. Records of the South 
Australian Museum 25(2): 1 13-120. (With D. P. 
Gordon as senior author). 

108. Discovery and identity of 110-year old Hutton 
Collection of South Australian Bryozoa. Records of 
the South Australian Museum 25(2): 121-128. 
(With D. P. Gordon as senior author). 



An aberrant new genus and subfamily of the spiculate 
bryozoan family Thalamoporellidae epiphytic on 
Posidonia. Journal of Natural History 25(5): 1363- 
1378. (With D. P. Gordon as senior author). 



1992 



110. A new genus of the bryozoan superfamily 
Schizoporelloidea, with remarks on the validity of the 
family Lacernidae Jullien, 1888. Records of the 
South Australian Museum 26(1): 67-71. (With D. P. 
Gordon as junior author). 

111. Review of the recent Scleractinia (stony corals) of 
South Australia, Victoria and Tasmania. Records of 
the South Australian Museum Monograph Series 
No. 3: 1-82. (With S. D. Cairns as senior author). 



64 



W. ZEIDLER 



In Press 

Hartmann-Schroder, G. & Parker, S. A. Four new species of 
the family Opheliidae (Polychaeta) from southern 
Australia. Records of the South Australian Museum 
28(1): 00-00. 

Hayward, P. J. & Parker, S. A. Notes on some species of 
Parasmittina Osburn, 1952 (Bryozoa: Cheilostomatida). 

Parker, S. A. & Cook, P. L. Records of the bryozoan family 
Selenariidae from Western Australia and South Australia, 
with the description of a new species of Seleharia Busk, 
1854. Records of the South Australian Museum. 27(1): 
1-11. 



Book Reviews 
1970 



1974 



1980 



Storr, G. M. 1973. 'List of Queensland Birds'. 
Reviewed in South Australian Ornithologist 26: 
146-148. 



Wagstaffe, R. 1978. Type Specimens of Birds in the 
Merseyside County Museums'. Reviewed in Emu 80: 
46. 

Etchecopas, R. D. & Hue, F. 1978. 'Les Oiseaux de 
Chine, de Mongolie et de Coree: Non Passereaux'. 
Reviewed in Emu 80: 174. 

Cheng Tso-Hsin. 1976. 'Distributional List of 
Chinese Birds'. Edn. 2. Reviewed in Emu 80: 174. 



References 



Mees, G. F. 1957-1969. 'A Systematic Review of the 
Indo-Australian Zosteropidae'. Parts 1-3. Reviewed 
in Emu 70: 94-95. 



1970-1978 



1971 



Ali, S. & Ripley, S. D. 1968-1974. 'Handbook of the 
Birds of India and Pakistan'. Vols 1-10. Reviewed in 
Emu 70: 37; 71: 144; 72: 119; 73: 141-142; 74: 
107-109, 270-271; 78: 166. 



McGill, A. R. 1970. 'Australian Warblers'. Reviewed 
in Emu 71: 90-91. 



EMERSON, K. C. & PRICE, R. D. 1986. Two new species 
of Mallophaga (Philopteridae) from the mallee fowl 
(Galliformes: Megapodiidae) in Australia. Journal of 
Medical Entomology 23: 353-355. 

HORTON, P. 1993. Obituary: Shane Alwyne Parker. South 
Australian Ornithologist 31: 148-150. 

PARKES, K. C. 1980. A new subspecies of spiny-cheeked 
honeyeater Acanthagenys rufogularis, with notes on 
generic relationships. Bulletin of the British 
Ornithologists' Club 100: 143-147. 

PRICE, R. D. & EMERSON, K. C. 1984. A new species of 
Megapodiella (Mallophaga: Philopteridae) from the 
mallee fowl of Australia. The Florida Entomologist 67: 
160-163. 



W. ZEIDLER, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Rec. S. Aust. Mus. 27(1): 57-64, 
1994. 



SOUTH 

AUSTRALIAN 

MUSEUM 



VOLUME 27 PART 1 
MAY 1993 
ISSN 0376-2750 



CONTENTS: 



ARTICLES 

1 S. A. PARKER & P. L. COOK 

Records of the bryozoan family Selenanidae from Western Australia and South Australia, 
with the description of a new species of Selenaria Busk, 1854 

13 DP. BARTON 

A checklist of helminth parasites of Australian Amphibia 

3 1 P. SUTTON 

Material culture traditions of the Wik people, Cape York Peninsula 

53 M.ZBIK&A. PRING 

The Chooikooning 001 meteorite: a new (L6) olivine-hypersthene chondrite from South 
Australia 

57 W. ZEIDLER 

Obituary - Shane A. Parker 



Published by the South Australian Museum, 
North Terrace, Adelaide, South Australia 5000. 



SOUTH 

AUSTRALIAN 

MUSEUM 



,994 



PROCEEDINGS OF THE FOURTH CONFERENCE ON AUSTRALASIAN 
VERTEBRATE EVOLUTION, PALAEONTOLOGY AND SYSTEMATICS, 

ADELAIDE, 19-21 APRIL 1994. 



INTRODUCTION 

During the third Conference on Australasian Vertebrate Evolution, 
Palaeontology and Systematics, held in Alice Springs in 1991, delegates were 
asked to decide the venue of the next meetings two years thence. Rhys Walkley 
had reminded me that 1993 would be the centenary of the first expedition to 
Lake Callabonna. It therefore seemed fitting that Adelaide should host the 1993 
meetings, and that in fact they should coincide with an exhibition about the 
Lake Callabonna fossils to be held at the South Australian Museum. 

This offer was accepted, together with a non-exclusive theme of Plio- 
Pleistocene palaeontology, with Rod Wells (Flinders University of South 
Australia) and Neville Pledge (South Australian Museum) combining to 
organise the meetings and pre- and post-conference excursions. The organisers 
are grateful for the help given by Mr Gavin Prideaux in the program planning 
and printing of the abstracts. 

The Conference was held in the Grosvenor Hotel, North Terrace, Adelaide, 
between 19-21 April 1993. More than fifty professionals, students and 
amateurs, several from overseas, attended to hear 49 papers read and to see 
eleven poster presentations. The student prize of an airline ticket to any 
Australian city for the purpose of study was donated by Australian Airlines. It 
was awarded to Ms Sanja Van Huet (Monash University) for her description of 
the Lancefield megafaunal site. 

A five-day pre-conference excursion to Lake Callabonna and Lake 
Palankarinna drew 13 participants, and an ad-hoc excursion to Naracoorte had 
20 people. 

During the Conference the exhibition 'Fossils of the Lake' was officially 
opened by Dr Richard Tedford (American Museum of Natural History), a 
veteran of two major expeditions to Lake Callabonna. The author is grateful to 
all who helped make the Conference a success. 



NEVILLE PLEDGE 

Curator of Fossils, 

South Australian Museum 



FOSSILS OF THE LAKE : A HISTORY OF THE LAKE CALLABONNA 

EXCAVATIONS 

Nevilles. Pledge 



Summary 

The discovery of Diprotodon in Lake Mulligan in 1892 led to a major expedition by the South 
Australian Museum the following year. Initial reports indicated scores, if not hundreds, of skeletons 
exposed on the dry lake bed. Excavation showed the reality: most skeletons were incomplete and 
the bones badly badly eroded or rotten. Nevertheless, a large quantity of fossilised material was 
recovered, under trying circumstances, by the two-stage expedition, and returned to Adelaide. 
Preparation and repair of the bones took many years, and the first complete Diprotodon skeleton 
was unveiled in 1907. Other fossils includes a new species of giant bird, Genyornis newtoni, the 
giant wombat Phascolonus gigas (which proved the synonym of Sceparnondon) and several large 
extinct kangaroos. 



FOSSILS OF THE LAKE: A HISTORY OF THE 
LAKE CALLABONNA EXCAVATIONS 



NEVILLES. PLEDGE 



PLEDGE, N. S. 1994. Fossils of the Lake: a history of the Lake Callabonna excavations. Rec. S. 
Aust. Mus 27(2): 65-77. 

The discovery of Diprotodon fossils in Lake Mulligan in 1892 led to a major expedition by 
the South Australian Museum the following year. Initial reports indicated scores, if not 
hundreds, of skeletons exposed on the dry lake bed. Excavation showed the reality: most 
skeletons were incomplete and the bones badly eroded or rotten. Nevertheless, a large quantity 
of fossilised material was recovered, under trying circumstances, by the two-stage expedition, 
and returned to Adelaide. Preparation and repair of the bones took many years, and the first 
complete Diprotodon skeleton was unveiled in 1907. Other fossils included a new species of 
giant bird, Genyornis newtoni, the giant wombat Phascolonus gigas (which proved the 
synonymy of See pamodori) and several large extinct kangaroos. 

Subsequent expeditions by R. A. Stirton in 1953 and R. H. Tedford in 1970 have thrown 
more light on this, Australia's first Fossil Reserve, a 'veritable necropolis'. 

N. S. Pledge, South Australian Museum, Adelaide, South Australia 5000. Manuscript received 
1 February, 1994. 



In 1831, while exploring the Wellington Valley 
of New South Wales, Major Thomas Mitchell 
found a fragment of a large jaw in one of the caves 
(Mitchell 1839). He sent it to Professor Richard 
Owen in London for identification. Owen, the 
world's leading anatomist of the day, soon realised 
that it came from a gigantic new species of 
marsupial, which he named Diprotodon, for its 
pair of large protruding incisor teeth ('two forward 
teeth') (Owen 1839). 

Over the next few decades, Owen described 
more and more bones of the skeleton of 
Diprotodon sent to him by collectors in Australia, 
and eventually (Owen 1870, 1877), he published 
a skeletal reconstruction of the animal. But he was 
continually frustrated by the lack of associated 
footbones and he died in December 1892 without 
knowing their complex structure, only months 
before discoveries of complete skeletons would be 
made. 

In 1870, a reward of £1 000 was offered for the 
discovery of a complete Diprotodon skeleton, and 
in the 1880s the discovery of skulls at Gawler 
(Howchin 1891) and Baldina Creek near Burra 
(Zietz 1890a), and associated bones at Bundey on 
the Murray Plains (Zietz 1890b), raised hopes that 
success would be attained. When word reached 
Adelaide in November 1892 that Diprotodon 
bones had been found on 'Mulligans Lake', north 
of Lake Frome, excitement swept the colony. 



Geomorphology and geology 

Originally called Lake Mulligan, Lake 
Callabonna is one of a chain of large dry saltpans 
(Fig. 1) that form a horseshoe around the northern 
Hinders Ranges from Lake Torrens to Lake Frome 
(Stirling 1894). 

On the site of an ancient silted-up lake system 
(Callen & Tedford 1978), modern Lake 
Callabonna has been formed by deflation: the 
wind blowing dried salt, sand and clay particles to 
the east and north to form the Strzelecki Desert. In 
earlier times, the lake was considerably larger and 
intermittently full of fresh water (Tedford 1993). 
Stranded beaches may be found kilometres from 
the present shoreline. 

During the Pleistocene, when the Diprotodon- 
bearing Millyera Formation was deposited, there 
was a fluctuating water level, as indicated by 
cyclic couplets of sand and laminated clay, the 
sand often being ripple-marked (shallow water or 
dry conditions) or desiccation cracked (dry/arid 
conditions). But the absence of fish fossils 
suggests the water in which the clay was 
deposited was too shallow or ephemeral to support 
them. Pieces of wood, and cones of the native pine 
Callitris indicate a wooded shoreline (Tedford 
1973). Today it is virtually bare. 

Later a more permanent lake formed supporting 
a plentiful fish and mollusc population. The 



66 



N. S. PLEDGE 




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FIGURE 1. View of Lake Callabonna, looking south from the hill behind the 1893 camp. Photo: attrib. H. Hurst. SA 
Museum Heritage Collection. 



mound springs then formed islands in the lake 
and were used as safe nesting areas for waterbirds 
whose bones have been found, and dated between 
2 000 and 3 000 years old. This may still occur 
today, such as during the floods of the early 1970s 
when the lake was full for several years. There is 
evidence that Aborigines sometimes visited these 
islands in search of food such as swans' eggs, 
since worked quartzite flakes have been seen on 
one of them, associated with bird bones and 
eggshell fragments (pers. observ.). 



Human history of the area 

Lake Callabonna falls within the sphere of the 
Pilatapa (Piladappa) Aborigines (Fig. 2) who 
occupied an area from Freeling Heights and Lake 
Frome to Lake Blanche and the southern part of 
the Strzelecki Desert, where an occupation site 
over 14 000 years old is known (Smith et al. 
1991). Stone artefacts, including grinding stones 
possibly obtained from Prism Hill, south of 
Balcanoona, and polished stone axes (possibly 
traded from the east) have been found near the 
lake (pers. observ.). Ancient rock engravings are 
known in the Flinders Ranges. It is possible that 
the legendary *Yamuti (Tunbridge 1988) of the 
nearby Adnyamathanha people is a folk memory 



of Diprotodon with which they could have been 
briefly contemporaneous, or at least a construction 
built on the knowledge of giant bones in the lake. 

Robert Stukey named it Mullachon Lake in 
1860, after the Aboriginal name for the mound 
springs on its western side. This soon became 
corrupted to Lake Mulligan. 

The first European to see Lake Callabonna was 
Edward John Eyre in 1840. A few years later 
(1844), the lake was visited by Charles Sturt's 
party coming in from the east. 

Pastoral activity started rather late, in the 1860s 
at Blanchewater. In 1 882, John Ragless 
established his son Frederick B. (Fig. 3) on the 
Callabonna run, on the eastern side of the lake 
(Mincham 1967). In establishing the homestead 
(Fig. 4) Frederick Ragless found Diprotodon 
bones in a well nearby and these were sent to the 
South Australian Museum in 1885. 



Discovery 

The Aborigines had long talked of large bones 
on the bed of Lake Mulligan, but they had been 
thought to be of bogged horses or cattle (Ragless 
1893, South Australian Register, 7 July 1893). 
On 12 June, 1892 Ragless had accompanied an 
Aboriginal stockman to an area on the lake bed to 



LAKE CALLABONNA EXCAVATIONS 



67 




FIGURE 2. Map of Lake Callabonna and environs, showing the Pilatapa tribal area (after Tindale 1974) and some 
pastoral boundaries. 



68 



N. S. PLEDGE 











mm. 



ill 

ill! 



II- 



FIGURE 3. 

Ragless. 



Frederick B. Ragless. Photo: courtesy L. 



see the bones (Fig. 5), following a conversation 
with the Aborigine a few days earlier when they 
had been talking about elephants. In his journal 
Ragless recorded that the boy, Jacky Nolan, had 
told him: 

I bin tinkit elephant alonga this country one time. Me 
bin see'em bones. Me show'em you big fella bone 
alonga lake. 'Im much too big alonga panto [horse] 
(Ragless, unpubl.) 

Ragless collected some of the bones and took 
them back to the homestead to show his men, and 
planned to take them to Adelaide at first 
opportunity. One of the men, John Meldrum, 
persuaded Nolan to take him back to the fossil site 
where he made his own collection. Shortly 
afterwards Meldrum 'left the station and came to 
town, where he exhibited the specimens at Mr P. 
Lee's Black Swan Hotel, North Terrace and 
claimed to be the discoverer' (South Australian 
Register, Thursday 6 July 1893, p. 4). Discovery 
of the bones led to a dispute over the reward which 
was never really settled. 

Meldrum' s arrival in Adelaide (South 
Australian Register, Tuesday 4 November 1892) 
with bones from Lake Callabonna was seized 
upon by the local press who followed succeeding 
events avidly. The Register (2 December 1892, p. 
5) reported that the South Australian Museum was 



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FIGURE 4. Callabonna Homestead, 1884. Photo: courtesy L. Ragless. 



LAKE CALLABONNA EXCAVATIONS 



69 



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RGURE 5. Diprotodon skeleton eroding out of the lake 
floor. Photo: attrib. H. Hurst. SA Museum Heritage 
Collection. 



sending Henry Hurst to investigate the site. He left 
on 2 December 1892 and returned on 5 January 
with great news. 



1893, Phase 1 

Hurst's favourable report to the Museum Board 
was enough for them to send him back with a 
larger expedition and more detailed instructions. 

Henry Hurst had worked for the Queensland 



Geological Survey and had collected fossils of 
Diprotodon etc. from the black soils of the Darling 
Downs, so he seemed well suited to lead the 
expedition which left on 13 January for three 
months to conduct systematic searches and 
excavations for Diprotodon specimens. With him 
he took his brother George, John Meldrum, two 
labourers, and an Afghan to handle the camels 
and bring supplies. The party travelled by rail as 
far as Farina (409 miles/658 km), where they 
obtained camels and horse and buggy (Fig. 6) to 
go around the north end of the Flinders to Lake 
Mulligan/Callabonna. This last stage of 200 miles 
(320 km) took eight days to travel. 

Hurst set up camp on an 'island' a couple 
kilometres from the edge of the lake bed, but after 
three weeks subject to the full force of the 
prevailing winds, moved it permanently to the 
western side of this island (Fig. 7). Excavations 
started at the spot first pointed out by Meldrum, 
but these specimens proved to be much decayed. 
Hurst used them, however, to instruct and train 
his inexperienced workers. 

Hurst's reports to Stirling were initially sent 
every few weeks and were extravagant in their 
description of the deposit and his results. Later 
they dealt more with the problems encountered - 
irregular and late mails and shortage of feed and 
stores (Figs 8, 9) due to rains, requirements for 



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FIGURE 6. Hurst's party about to leave Bruce' s Hotel, Farina, with camel team and buckboard, 1893. Photo: attrib. 
H. Hurst. SA Museum Heritage Collection. 



70 



N. S. PLEDGE 




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FIGURE 7. Permanent camp. Lake Mulligan 1893. Photo: attrib. H. Hurst. SA Museum Heritage Collection. 





-*.- 



FIGURES 8,9. All supplies were brought in by camel 
from many kilometres away. 8, water; 9, firewood. 
Photos: attrib. H. Hurst. SA Museum Heritage Collection. 



sending the bones back to Adelaide and 
impending deadlines for the return of the camels 
and the cessation of the expedition. Hurst did, 
however, send accurate reports of new discoveries, 
such as the footpad impressions associated with 
the Diprotodon foot bones (letter, 16 March 
1893), and the partial skeleton of a giant 
'struthious' bird (letter, 6 May 1893), later to be 
named Genyornis newtoni. His excitement about 
the latter was such that he telegraphed the news of 
its discovery to Stirling. There was also a 
fossilised fruit, found near the bird remains, which 
he sent to Adelaide for identification. According 
to Hurst, Professor Ralph Tate (Adelaide 
University), identified it as Frenela (letter from 
Hurst, 20 June 1893), the old name for Callitris 
referred to by Stirling (1900). 

Hurst suggested that he collaborate with Stirling 
in describing the Diprotodon remains, took a 
number of photographs of the operations (Figs 10, 
1 1) and invited Stirling to see the site for himself. 
Occasional rain showers impeded work but by 25 
May when the Government Geologist, H. Y. L. 
Brown, visited the site, 'three nearly complete 
skeletons of the diprotodon... [had] been unearthed 
(Fig. 12), together with over 2000 separate bones 
of the same animal belonging to over seventy 
individuals' (South Australian Register 23 May, 
1893, p. 5). Stirling conveyed this information to 



LAKE CALLABONNA EXCAVATIONS 



71 



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FIGURE 10. Excavating. The posted sign registers the area as a mining 'claim', but the details are illegible. The men 
are not identified but could include Meldrum. Photo: attrib. H. Hurst. SA Museum Heritage Collection. 



the scientific world in a letter published in The 
Times (28 April 1893). 

Brown's report (South Australian Register, 1 
July 1893, p. 6; 1894) generally supported Hurst's 
claims of the fossil deposit. He noted that 
articulated footbones, encased in limy concretions 
that preserved apparent impressions of the sole of 
the foot, had been found. He further noted that 
bones of a giant wombat, kangaroos and a 
gigantic bird occurred in the deposit and he 
concluded by recommending 'that the whole area 
of the lake be reserved' for future scientific 
exploration, 'to prevent the indiscriminate digging 
up and removal of portions of the specimens.' This 
sentiment was heartily endorsed by the South 
Australian Register (31 July 1893, p. 6). 

When heavy rain came in July, Hurst and 
Meldrum returned to Adelaide with some of the 
many bones they had excavated, leaving the rest 
of the party to continue work. Hurst's verbal report 
to the Museum Committee members enthused 
them enough to appropriate another £100 to cover 
his expenses. In addition two drays were to be 
despatched from Hawker to bring all the 
remaining bones from Lake Callabonna at an 



estimated cost of a further £100. Hurst was to 
return to the lake forthwith to supervise this 
packing and dispatch, and await further 
instructions. 

While in Adelaide, Hurst completed (29 July 
1893) a preliminary report to the Museum 
Committee on his journey, excavations and 
geological observations. This formed the basis for 
Stirling's subsequent articles (Stirling 1900). 
Hurst also received a letter from his brother 
George, still at Lake Callabonna, reporting 
excitedly: 

As soon as T had dug under the pelvis and into the 
exact spot where the pouch would be, I came on a 
dear little diprotodon humerus about six inches long. 
It is evident the large animal is a female and had a 
piccaninny diprotodon in her pouch when she died... 
The claim is very wet and the bones are very difficult 
to remove but you can depend I will get them all as I 
think this is the most wonderful discovery ever made 
in the world. (George Hurst July 1893, fide Tcdford 
1973). 

This discovery exemplifies a problem that was 
beginning to emerge, for while some of these 
neonate bones have survived, they were, either at 



72 



N. S. PLEDGE 




FIGURE 11. Detail of the specimen in Fig. 10. Photo: attrib. H. Hurst. SA Museum Heritage Collection. 



Ihe time of excavation or subsequently in 
Adelaide, separated from those of the putative 
mother. Stirling was apparently already having 
misgivings about the procedures being used by 
Hurst, who had been concentrating on collecting 
bones that until then were unrepresented or poorly 
known in collections, and only rarely keeping 
associated bones together. Had the bones of 
mother and baby been kept together, the confusion 
over whether the two forms at Lake Callabonna 
were different sexes or different species (e.g. 
Williams 1982: Appendix IV) might have been 
resolved before now. 

Stirling and his assistant, A. H. C. Zietz (Figs 
13, 14), spent considerable time sorting through 
the bones brought down by Hurst, arranging them 
anatomically. Many had never been seen in such 
perfect condition, and those of the feet, tail and 
pouch (epipubes) had not been seen before (South 
Australian Register 31 July 1893, p. 6). Many 
had suffered during their journey to Adelaide 
because of poor packing though, and there was 
additional disappointment that no complete 
skeleton had been collected. 

Stirling hoped that three or four complete 



skeletons would be sent down to Adelaide in time 
to be exhibited at the September meeting of the 
Australasian Science Association (South 
Australian Register 1 August 1893, p. 5). 

Hurst went back to the lake early in August, 
and was followed on 1 1 August by Stirling, Zietz 
and Thomas Cornock, the Museum's taxidermist 
and articulator. 

1893, Phase 2 

After inspection of the camp and excavation 
site, Stirling 'was very seriously dissatisfied with 
the way in which the work had been and was 
being carried on, [and] gave Mr Hurst and his 
brother a week's notice of the termination of their 
engagement, on which they with his consent, left 
at once.' The other men stayed on. It rained during 
Stirling's stay and he was unable to carry out the 
exploration he had hoped to do. After reorganising 
the party, he put Zietz in charge, confident of the 
work being done satisfactorily, and left for 
Adelaide on 21 August (Minutes, special 
committee meeting 31 August 1893). 

Thomas Cornock, who had accompanied 



LAKE CALLABONNA EXCAVATIONS 



73 




FIGURE 12. Partially excavated Diprotodon skeleton. This specimen is lying on its belly, head to the right. Photo: 
attrib. H. Hurst. SA Museum Heritage Collection. 



Stirling and Governor Kintore to Darwin 
(Palmerston) in 1891, made a number of pencil 
sketches of specimens and the excavations, which 
have only recently been rediscovered. 

Following Hurst's departure and Zietz's 
assumption of control of the excavations, there 
seems to have been little news from the field until 
Zietz's return and his interview by a reporter from 
the Adelaide Observer (9 December, 1893, p. 33). 
Hurst had filled twenty-eight wooden cases with 
bones. Zietz added nearly sixty more. There was 
one complete skeleton of Diprotodon, in good 
condition, another not so good, and a third of 
possibly another, smaller species. Amongst other 
things, they had collected one skull of the giant 
bird and fragments of another one, a skeleton of 
the giant wombat and two skeletons of extinct 
kangaroos, as well as a second juvenile 
Diprotodon (Fig. 15). 

Weather and working conditions deteriorated as 
the year wore on. The drought worsened, as did 
the rabbit plague. Constant dust storms caused 
'sandy blight* eye problems, and the dying rabbits 
and bad water contributed to gastric illness. 
Finally heavy rains in November made excavation 



impossible and the party returned to Adelaide 
amid great interest from the public and praise 
from Stirling. 

The expedition had originally been financed by 
the Museum - £250 being scraped from their 
meagre budget for three months work. This soon 
ran out and the operation was saved by a generous 
gift of £500 from Sir Thomas Elder. The Museum 
was also indebted to the Surveyor General, Mr G. 
W. Goyder and Mr Peter Waite for services such 
as supplying camels. Mr Frederick Ragless 
assisted in various practical ways. Ragless and 
Meldrum both applied for the reward for the 
discovery of the Diprotodon skeletons. The 
Museum Board rejected Meldrum's claim in 
favour of Ragless, but because of the huge 
financial costs engendered by the expeditions, they 
were unable to pay the reward. 



The specimens 

Despite the various problems encountered by 
the 1893 expeditions, a large number of bones 
were collected. By far the most numerous were the 



74 



N. S. PLEDGE 







-1 



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I 



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FIGURE 13. Dr Edward C. Stirling. Photo: SAM Photo 
Library. 



FIGURE 14. A. H. C. Zietz. Photo: SAM Photo Library. 



remains of Diprotodon - some eighty specimens 
were investigated and at least partially collected. 
Also collected were partial skeletons of several 
giant birds (Genyornis), two giant wombats 
(Phascolonus) and some kangaroos (Sthenurus 
and Macropus), as well as the fragmented 
impressions of Diprotodon foot pads, and a mass 
of comminuted twigs found in the gut region of a 
Diprotodon (Stirling & Zietz 1899). 

Reading through Hurst's letters, it is apparent 
that the list should be larger than this. For 
instance, he mentions (20 June 1893) several 
skeletons of a new wombat just found by his 
brother, that may have 'exceeded a bullock in 
size', and 'a minute mammal which must prove 
new to science. The tibia of the latter measures 
about 2V2 inches [6.3 cm] in length'. Although 
Hurst intended bringing the latter with him to 
Adelaide, Stirling apparently never saw it and its 
whereabouts (and identity) is unknown. 



Creation of the fossil reserve 

The South Australian Government Geologist H. 
Y. L. Brown visited Lake Callabonna in June 



1893. At the end of his report (1894) he 
recommended that the lake bed be proclaimed a 
Fossil Reserve 'to prevent the indiscriminate 
digging up and removal of portions of the 
specimens' (the suggestion came at a time when 
an Adelaide syndicate was being floated to do just 
that). 

Brown's far-sighted suggestion was duly 
followed, and on 5 December 1901 the Lake 
Callabonna Fossil Reserve was gazetted. It is 
believed to be the first fossil locality in Australia, 
and one of the first in the world, to be so declared. 



Aftermath 

Following his return to Adelaide with the bulk 
of the fossils found at Lake Callabonna, Zietz 
immediately plunged into the task of sorting and 
repairing bones, and compiling a composite set 
from which to cast a complete skeleton (Stirling 
1907). This took many years. The problem was 
that the bone was impregnated with salts, mainly 
sodium sulphate, which soaked up moisture 
during winter and made cleaning and restoration, 
with the facilities, methods and materials then 



LAKE CALLABONNA EXCAVATIONS 



75 




: 




4$ 




FIGURE 15. Associated bones of the juvenile Diprotodon found by Zietz's party, August 1893. SAM P10563 Photo: 
N. Pledge. 



available, very difficult (Hale 1956). Meanwhile, 
he assisted Stirling in describing the giant bird 
Genyornis, and the feet of Diprotodon (Stirling & 
Zietz 1896, 1899a,b). Casting of the bones was 
done by Robert Limb, father of the entertainer of 
the same name. A restoration painting of 
Diprotodon in life was done by C. H. Angas 
(Stirling 1907). 

For his part, Stirling decided in 1894 that the 
lake should be called Lake Callabonna (Board 
Minutes, 2 February 1894) after Ragless' sheep 
run that bordered it on the east. It had never 
formally been named - its common name, Lake 
Mulligan, was a corruption of 'Mullachon', the 
name given it in 1860 by Robert Stukey. This was 
the Aboriginal name for the mound springs on the 
western side that still are known as Mulligan 
Springs. 

Stirling's study of the fossil remains resulted in 
several short articles (1893a,b, 1894a,b, 1896a,b, 
1900, 1907), and a series of six monographs 
published in the specially created Memoirs of the 
Royal Society of South Australia between 1899 
and 1913. 

While Zietz was working on the skeleton of 
Diprotodon, Stirling travelled overseas, visiting 
museums. Several were interested in obtaining a 
cast or duplicate specimens of bones of 



Diprotodon, and in this way he obtained by 
exchange the dinosaur Diplodocus leg bones on 
display on Level 1 of the South Australian 
Museum. Seven casts of the Diprotodon skeleton 
were eventually distributed. 

Preparation of the Diprotodon cast was 
completed in 1906 and unveiled to the public in 
1907, together with numerous other specimens. 



Conclusions 

After 1893, Lake Callabonna lay almost 
forgotten and undisturbed by scientists for sixty 
years. The only known visits were by Robert 
Bedford, of the privately-owned Kyancutta 
Museum, in 1928 (Cooper 1987) and H.O. 
Fletcher (1948) from the Australian Museum. It is 
not known what Bedford collected, but Fletcher 
found nothing, apparently having gone to a 
different part of the lake, away from the 1893 site. 

In 1953 the Board of the South Australian 
Museum, at the urging of new member C. W. 
Bonython, suggested to University of California 
researcher Professor R. A. Stirton, visiting 
Australia in search of Tertiary mammals, that 
reinvestigation of Lake Callabonna would at least 
bring some positive results for his expedition. 



76 



N. S. PLEDGE 



Stirton and Fulbright student R. H. Tedford, 
together with Museum and University of Adelaide 
personnel (Stirton 1954) spent three weeks 
excavating near the site of the 1893 camp which 
they were able to relocate. Several specimens of 
Diprotodon were collected and taken back to the 
University of California Museum of Paleontology 
at Berkeley to be prepared and ultimately studied. 

Tedford became fascinated by the locality and 
in 1970 led a joint expedition with the 
Smithsonian Institution and South Australian 
Museum, with excellent results (Tedford 1973). 
Although Diprotodon was the first objective, more 
important specimens were of Genyomis (e.g. Rich 
1979), Phascolonus, Protemnodon and Sthenurus 
(Wells and Tedford in prep.). In 1983 T. Rich and 
P. V. Rich, with logistic support from the 
Australian Army, returned to search (fruitlessly) 
for more Genyomis material. 

Stirton' s 1953 trip could be said to be the 
beginning of modern mammalian palaeontology 
in Australia, since it stirred academic and public 
interest and spawned a growing number of 



Australian and American students with an interest 
in the Tertiary and Quaternary of this continent. 

We must not, however, forget the thoughtful 
and interested pastoralists such as Frederick 
Ragless and enthusiasts like Henry Hurst, and 
explorers such as Thomas Mitchell who initially 
brought these fascinating fossils to scientific and 
public notice. 

Acknowledgments 

I am indebted to Ms J. M. Scrymgour and her long- 
time interest in this story for accumulating much of the 
information and photographs used herein. I thank also 
Mr E. S. Booth, grandson of Sir Edward Stirling, and Mr 
Leigh Ragless, grandson of Frederick Ragless, for their 
interest in providing family photos or records for my use 
in the exhibition 'Fossils of the Lake' (South Australian 
Museum, 19 April - 22 August 1993) and subsequently 
in this article. Ms Jenni Thurmer supplied original glass 
plate negatives from the South Australian Museum's 
Heritage Collection. Mr Philip Jones provided assistance 
with archival research. The manuscript was typed by Ms 
Debbie Churches. 



References 



BROWN, H. Y. L. 1894. Annual Report of the 
Government Geologist, for the year ended June 30th, 
1894. Government Printer, Adelaide. 

CALLEN, R. A. & TEDFORD, R. H. 1976. New late 
Cainozoic rock units and depositional environments, 
Lake Frome area, South Australia. Transactions of 
the Royal Society of South Australia. 100(3): 125- 
167. 

COOPER, B. J. 1987. Historical Perspective: the 
Kyancutta Museum. Quarterly Geological Notes. 
Geological Survey of South Australia. 103: 2-3. 

FLETCHER, H. O. 1948. Fossil Hunting 'West of the 
Darling' and a visit to Lake Callabonna. Australian 
Museum Magazine 9(9): 315-321. 

HALE, H. 1956. The First Hundred Years of the South 
Australian Museum 1856-1956. Records of the South 
Australian Museum. 12: 1-225. 

HOWCHIN, W. 1891. (Abstract of Proceedings). 
Transactions of the Royal Society of South Australia. 
14:361. 

MINCHAM, H. 1967. Vanished giants of Australia. 
Rigby, Adelaide. 

MITCHELL, T. L. 1839. 'Three Expeditions into the 
Interior of Eastern Australia; with description of the 
recently explored region of Australia Felix, and of the 
present colony of New South Wales.' Two volumes. 
T. & W. Boone, London. 

OWEN, R. 1839. Letter, in MITCHELL, T. L. Three 
expeditions ... Vol. 2, pp. 365-369. 



OWEN, R. 1870. On the fossil mammals of Australia- 
Part III. Diprotodon australis, Owen. Philosophical 
Transactions of the Royal Society, London 160: 519- 
578, pis 35-50. Reprinted (1877) in 'Researches on 
the fossil remains of the extinct mammals of Australia 
with a notice of the extinct marsupials of England'. 

RICH, P. V. 1979. The Dromornithidae. Bureau of 
Mineral Resources Bulletin 184: 1-194. 

SMITH, M. A., WILLIAMS, E. & WASSON, R. J. 1991 . 
The archaeology of the JSN site: some implications 
for the dynamics of human occupation in the 
Strzelecki Desert during the late Pleistocene. Records 
of the South Australian Museum. 25(2): 175—192. 

STIRLING, E. C. 1893. [Extract from a letter concerning 
the discovery of Diprotodon and other mammalian 
remains in South Australia]. Proceedings of the 
Zoological Society, London 1893: 473-475. 

STIRLING, E. C. 1894. The recent discovery of fossil 
remains at Lake Callabonna, South Australia. Nature 
(London) 50: 184-188, 206-21 1, figs. 1-3. 

STIRLING, E. C. 1894. Supposed new extinct Gigantic 
Bird in Australia. Ibis series (6)6: 328. 

STIRLING, E. C. 1894. The new extinct Gigantic Bird 
of Australia. Ibis series (6)6: 577-578. 

STIRLING, E. C. 1896. The new extinct Gigantic Bird 

of Australia. Ibis series (7)2: 430. 

STIRLING, E. C. 1896. The newly discovered extinct 
gigantic bird of South Australia. Ibis series (7)2: 593. 



LAKE CALLABONNA EXCAVATIONS 



77 



STIRLING, E. C. 1900. Physical features of Lake 
Callabonna. Memoirs of the Royal Society of South 
Australia., I, i-xv, pi. A. Abstr., Nature (London) 61: 

275-278, figs. 1,2. 

STIRLING, E. C. 1901. [Diprotodon australis.) 
Museum Journal. (London) 1: 114-115 (1901-1902) 

STIRLING, E. C. 1907. Report of the Museum Director. 
South Australia. Public Library, Museum and Art 
Gallery, Adelaide; Report of the Board of Governors, 
1906-07: 8-9. 

STIRLING, E. C. 1907. Reconstruction of Diprotodon 
from the Callabonna deposits, South Australia. Nature 
(London) 76: 543-544, figs. 1, 2. 

STIRLING, E. C. 1913. Fossil remains of Lake 
Callabonna. Part IV. 1. Description of some further 
remains of Genyornis newtoni, Stirling and Zietz. 2. 
On the identity of Phascolomys (Phascolonus) gigas, 
Owen, and Sceparnodon ramsayi, Owen, with a 
description of some of its remains. Memoirs of the 
Royal Society of South Australia 1: 111-178, pis. 
XXXV-LVII. 

STIRLING, E. C. & ZIETZ, A. H. C. 1896. Preliminary 
notes on Genyornis newtoni, a new genus and species 
of fossil struthious bird found at Lake Callabonna. 
Description of the bones of the leg and foot of 
Genyornis newtoni, a fossil struthious bird from Lake 
Callabonna. Transaction of the Royal Society of 
South Australia 20: 171-211, pis. III-V. 

STIRLING, E. C. & ZIETZ, A. H. C. 1899. Preliminary 

notes on Phascolonus gigas, Owen (Phascolomys 
[Phascolonus] gigas, Owen). Transactions of the 
Royal Society of South Australia 23: 123-135. 

STIRLING, E. C. & ZIETZ, A. H. C. 1899. Fossil 
remains of Lake Callabonna. Part I. Description of the 
manus and pes of Diprotodon australis, Owen. 
Memoirs of the Royal Society of South Australia 1: 



1-40, pis. I-XVIII. Abstr., Nature (London) 61: 275- 
278, 2 figs. (1900). 

STIRLING, E. C. & ZEITZ, A. H. C. 1900. Fossil 
remains of Lake Callabonna. Part II. Genyornis 
newtoni, a new genus and species of fossil struthious 
bird. Memoirs of the Royal Society of South Australia 
1: 41-80, pis XIX-XXIV. (Cf. 1896a) Abstr., Nature 
(London), LXIII, 181-182. 

STIRLING, E. C. & ZEITZ, A. H. C. 1900. Fossil 
remains of Lake Callabonna. Part III. Description of 
the vertebrae of Genyornis newtoni. Memoirs of the 
Royal Society of South Australia 1: 81-110, pis. 
XXV-XXXV. 

STIRTON, R. A. 1954. Digging Down Under. Pacific 
Discovery 7(2): 2-13. 

TEDFORD, R. H. 1973. The Diprotodons of 
Callabonna. Australian National History. 17: 349- 
354. 

TEDFORD, R. H. 1994. Lake Callabonna: 'Veritable 
necropolis of gigantic extinct marsupials and birds'. 
CAVEPS Abstracts (this volume). Records of the 
South Australian Museum 27: 217—229. 

TINDALE, N. B. 1974. 'Aboriginal Tribes of Australia' . 
University of California Press: Berkeley. 

TUNBRIDGE, D. 1988. 'Flinders Ranges Dreaming'. 
Aboriginal Studies Press: Canberra. 

WILLIAMS, D. L. G. 1982. The late Pleiostocene of the 
Flinders and Mt. Lofty Ranges, S.A. Unpublished 
Ph.D. Thesis, Flinders university of South Australia, 
Adelaide. 

ZIETZ, A. H. C. 1890a. Diprotodon remains. 
Transactions of the Royal Society of South Australia. 
13: 236. 

ZIETZ, A. H. C. 1890b. (Abstract of Proceedings). 
Transactions of the Royal Society of South Australia. 
13: 245. 



SUCCESSION OF PLIOCENE THROUGH MEDIAL PLEISTOCENE 
MAMMAL FAUNAS OF SOUTHEASTERN AUSTRALIA 

Richard H. Tedford 



Summary 

Southeastern Australia contains the most completely documented Pliocene-Pleistocene faunal 
succession known on the continent. Magnetostratigraphic, geochronologic and palaeontologic data 
from the Murray and coastal Victorian basins provide a chronology for local faunas covering early 
Pliocene through medial Pleistocene time. Changes in the composition of these assemblages appear 
to be congruent with the physical and palaeobotanical history of southeastern Australia. 



SUCCESSION OF PLIOCENE THROUGH MEDIAL PLEISTOCENE MAMMAL FAUNAS 

OF SOUTHEASTERN AUSTRALIA 



RICHARD H. TEDFORD 



TEDFORD, R. H. 1994. Succession of Pliocene through Medial Pleistocence mammal faunas of 
Southeastern Australia. Rec. S. Aust. Mus. 27(2): 79-93. 

Southeastern Australia contains the most completely documented Pliocene-Pleistocene faunal 
succession known on the continent. Magnetostratigraphic, geochronologic and palaeontologic 
data from the Murray and coastal Victorian basins provide a chronology for local faunas 
covering early Pliocene through medial Pleistocene time. Changes in the composition of these 
assemblages appear to be congruent with the physical and palaeobotanical history of 
southeastern Australia. 

The early Pliocene marine transgression in the Murray and coastal basins was accompanied 
by the spread of rainforest containing diverse arboreal mammals and genera now restricted to 
the tropics. At the generic level early Pliocene assemblages resemble later faunas and include 
the earliest appearance of some living and Pleistocene genera along with genera known only 
from the early Pliocene. Marine regression in the medial Pliocene restored more continental 
climates to southeastern Australia but the presence of a large lake in the Murray Basin sustained 
wet sclerophyll vegetation. Living genera whose species now occupy inland environments began 
to appear there. 

At the close of the Pliocene and beginning of the Pleistocene, forms close to living species 
appear as do species of extinct genera that became common elements in medial and late 
Pleistocene faunas. Wet sclerophyll vegetation was restricted to the highlands and drier 
shrubland associations developed in the lowlands as the temperate latitudes of Australia 
responded to global climatic cooling. By the time more rapid cycling of world climate appears in 
the medial Pleistocene, the southeastern Australian fauna included most of the associations of 
taxa that characterized the later part of the Quaternary. 

Richard H. Tedford, American Museum of Natural History, New York, N. Y. 10024. 
Manuscript received 19 April 1993 

Data reviewed by Woodburne et al. (1985) on shifting habitat for some forms. The pygmy 

the succession of mammal faunas in the Murray possum Burramys is a case in point in which the 

and adjacent Otway and Gippsland basins of Pliocene species were associated with lowland 

southeastern Australia suggested their singular rainforest yet its living species is an inhabitant of 

potential in defining a Pliocene and Pleistocene temperate alpine environments (Flannery et al. 

biostratigraphy for that part of the Australian 1992). It is also important to remember that 

continent. The locations of most of these faunas knowledge of the succession of local faunas and 

are shown in Figure 1. Subsequent study of these their contained taxa is limited by the geological 

assemblages, and further calibration of their record of southeastern Australia. Therefore the 

succession using paleomagnetic methods (An et observed stratigraphic ranges in that area represent 

al. 1986; MacFadden et al. 1987; Whitelaw 1989, only parts of the total ranges of the taxa. Further 

1991, 1993) has made it possible to look in more perspective on the temporal ranges of some 

detail at the temporal ranges of taxa and to Pliocene taxa can be made by comparison with the 

correlate faunal change with physical and biotic Lake Eyre Basin and northern Queensland where 

events deduced from other data. similarly calibrated faunas are known. 

The purpose of this contribution is to examine As Whitelaw (1991) pointed out, the 

the chronologic ranges of taxa, principally at the magnetostratigraphies relating to individual local 

generic level, and then to consider the possible faunas cannot in themselves be correlated with the 

ecological factors that might have guided the Geomagnetic Polarity Time Scale without 

succession based on genera whose living species evidence from other geochrono logical disciplines, 

have known habitat preferences. Such His work on the faunas of southeastern Australia 

extrapolations must be tempered by evidence of emphasized those assemblages from deposits 



80 



R. H.TEDFORD 



140 




-3 2 S 



-38 




FIGURE l . Map of south-eastern Australia showing location of sites from 
which the local faunas mentioned in the text were obtained. Basin margins 
marked approximately by the 500 ft. contour, highlands above that 
indicated by shading. The local faunas are listed vertically in ascending 
stratigraphic order separated by solid lines where in direct superposition 
and dashed lines where not. 



interbedded with volcanic rocks that have been 
dated using radioisotopic methods. In the case of 
the Nelson Bay Local Fauna (MacFadden et at. 
1987; Whitelaw 1991) the presence in the deposits 
of age-diagnostic planktonic foraminifera provided 
additional calibration. The Murray Basin sites lie 
within a longer magnetostratigraphic section that 
An et al. (1986), and Whitelaw (1991) correlate 
as lying within the Gauss through early Matuyama 
chrons using the pattern of polarity reversals. In 
any case, the chronological data constrain the 
fossil assemblages only to spans of time; shorter 
where the data are best, and longer where more 
loosely defined. The correlation chart (Fig. 2) 
shows these estimated ranges, and the range chart 
(Fig. 3) uses these estimates of maximum and 
minimum temporal range for taxa. In addition the 



total known range is indicated both within and 
outside southeastern Australia. 

This paper follows Cande and Kent (1992) in 
the calibration of epoch and informally designated 
subepoch boundaries: Miocene-Pliocene 
boundary, 5.3 Ma; early Pliocene-medial Pliocene 
boundary is the Gilbert-Gauss boundary, 3.55 Ma; 
medial Pliocene-late Pliocene boundary is the 
Gauss-Matuyama boundary, 2.60 Ma; Pliocene- 
Pleistocene boundary, is at the end of the Olduvai 
Subchron, Matuyama Chron 1.76 Ma; early 
Pleistocene-medial Pleistocene boundary is 
informally set at the end of the Matuyama Chron, 
0.78 Ma; medial Pleistocene-late Pleistocene 
boundary is informally set at 0.40 Ma; late 
Pleistocene-Holocene boundary is at 0.01 Ma 
(Ma: million years on the geological time scale; 



PLIO-PLEISTOCENE MAMMALS OF SOUTHEASTERN AUSTRALIA 




2- 



4- 



5- 



GPTS 



OTWAY AND PORT PHILLIP BASINS 



LATE 

PLEISTOCENE 



MEDIAL 
PLEISTOCENE 




EARLY 
PLEISTOCENE 



LATE 
PLIOCENE 



MEDIAL 
PLIOCENE 



o 




U^G 



EARLY 
PLIOCENE 



MURRAY 
BASIN 




MIOCENE 



IP 






FIGURE 2. Approximate limits in time for the local faunas discussed in the text as deduced from palaeomagnetic 
data and correlations to the Geomagnetic Polarity Time Scale (GPTS). Although each local fauna is essentially a 
point in geologic time this cannot be more closely constrained than the limits shown (Whitelaw 1991). The Sunlands 
Local Fauna from the Murray Basin has no supporting paleomagnetic data, but is given a chronologic position 
correlative with assemblages in the Otway Basin that belong to the early phase of the Pliocene transgression recorded 
in both basins. Calibration of the GPTS follows Cande and Kent (1992). 



M.y.: span of time, millions of years long). 

Extinct taxa are indicated in the text by 
preceding asterisk *. 



The faunal record 

The geochronological range chart (Fig. 3) plots 
the time spans for selected genera of marsupials 
and one family of rodents. These are arranged to 
show the succession of forms, but the following 



analysis of this record will discuss each family in 
systematic order. The best available faunal lists 
are given by Rich et al. (1991 ) and the sources for 
these lists are given there. Since 1991, additional 
faunas, pertinent to this review, have appeared 
including those in Flannery et al. (1992); Pledge 
(1992); Tedford and Wells (1992) and Turnbull et 
al. (1992 and 1993). Whitelaw (1991, 1992) gives 
lists for the Pleistocene faunas and references. An 
asterisk indicates extinct taxa. Two sites deserve 
further discussion: 



82 



R. H.TEDFORD 



AGE 



TAXON Ns 


\ 5.0 4.0 


3.0 


2.0 


1-0 




Ma 




Ramsay ia 
Petrogale 














—M 




Z 


















CO 
< 


Osphranter 
















--m 




CO 


Lasiorhinus 
















--■ 




>- 


Lagos trophus 
















■ 




< 


Satanellus 


















NA 


3 


Dasyuroides 
Sthenurus 




NA 












H 


NA 


5 


















Phascolonus 
Macro pus 










9 






--H 












iganteus / t ita 


n 








Muridae 












im 


optatum 


-H 






Diprotodon 

Vombatus 

Sarcophilus 


_| UTS 


is 


/harrisj 









laniarius 








parryi 


Notamacropus 












Thylogale 




mm 






I bi 


color 




— ■ 




CO 

Z 


Wallabia 




Propleopus 




— - 












--H 




Dorcopsis 


















NG 


CO 

< 


Dendrolagus 


















NA,NG 


CO 


Hypsiprymnodon 


NA 




— — 


— — 




— 


pereg 


rir 


us 


— Hi 


NA 


< 

I— 


Pseudocheirus 






Strigocuscus 


NA 


















Trichosurus 
Thylacoleo 


NA 










— 


cart 


lift 


ix 


-~H 




co 

< . 
o 










Simosthenurus 




- — 








parvus 


--H 

ml 




U 


Palorchestes 
Protemnodon 







mmmm 


1 




anak 













— 


trilobus 


-M 


Zygoma t u ru s 






Troposodon 




mam 


mmm 












\ 




EA,NA 




Baringa 






Ektopodontidae 

Euowenia 

Kurrabi 


NA 


EA • • 








































E. PLIOCENE 


M. PLIO. 


L.PLIO. 


E.PLEIST. 


M.R 


L.P. 


^HOL 



FIGURE 3. Geochronologic ranges for selected taxa discussed in the text following the data of Fig. 2. Taxa divided 
into coastal Victoria and Murray basins with regard to first occurrences. Dashed connections of occurrences refer to 
southeastern Australia (including South Australia), dotted connections indicate earlier or later occurrences in other 
parts of the continent [EA, eastern Australia; NA, northern and central Australia, NG, New Guinea] for taxa whose 
ranges begin outside southeastern Australia and others that become restricted with time. 



PLIO-PLEISTOCENE MAMMALS OF SOUTHEASTERN AUSTRALIA 



83 



The newly described Curramulka Local Fauna 
(Pledge 1992) from Yorke Penninsula, South 
Australia (Fig. 1 ) appears on stage-in-evolution 
criteria of its marsupials to predate the 
assemblages discussed here. The fact that it 
contains several genera that are components of 
Pliocene and Pleistocene faunas of southeastern 
Australia suggests that the Curramulka Local 
Fauna is at least of late Miocene age. The local 
fauna contains a number of arboreal forms but 
lacks those having species in present-day 
rainforest. It provides valuable perspective in our 
assessment of the history of certain genera and 
relevant information from this fauna will be 
included in the following summaries and 
discussion. 

The Beaumaris marsupial assemblage from the 
Port Phillip Basin (Fig. 1) consists of five 
specimens collected over the past one hundred 
years from outcrops of the Black Rock Sand 
Member of the Sandringham Formation at 
Beaumaris. Producing strata include the nodule 
bed that lies directly on a surface cut on the medial 
Miocene Fyansford Formation, and the overlying 
calcareous and ferruginous sands. None of the 
material has precise provenance, but differences in 
permineralization suggest different geochemical 
environments that may be indicative of 
stratigraphic position. In an effort to test the 
provenance of the material Gill (1957a) 
determined the flourine-phosphate ratios of the 
bony parts of three of the specimens and found a 
spread of values separated by only 0.5-0.6%. 
Similar determinations from the bony parts of 
shark teeth, some known to be from the ''nodule 
bed," produced a similar range of values. Gill 
concluded that the nodule bed was the probable 
source of the marsupial bones. Despite this there 
is still the possibility that the Beaumaris 
marsupial assemblage is a composite of older 
remanie and younger autochthonous material. 
Carter (1978) examined foraminifera from the 
"nodule bed" and found remanie material, the 
youngest of which was of medial Miocene age. 
Miocene squalodont whale teeth and the nautiloid 
*Aturia from Beaumaris also suggest that they are 
remanie lying within the "nodule bed" on the 
surface of disconformity. 

The heavily permineralized *Kolopsis sp. right 
ramus (Nat. Mus. Vict. P15911) described by 
Stirton (1957) and Woodburne (1969 as 
Zygomaturus gilli) is water-rolled and bored from 
exposure on the marine platform developed on the 
Fyansford Formation. A left ramus of *Kolopsis 
sp. (Nat. Mus. Vict. PI 6279), purchased by the 



Museum of Victoria from Albin Bishop in 1910 
and thought to be from Chinchilla, Queensland 
(Stirton 1957: 128), may actually be from 
Beaumaris because it is encrusted with calcareous 
tubes of spirorbid worms (T. Rich, pers. com. 
1979) and preserved in a manner similar to NMV 
P15911. These specimens may be remanie. 
Murray et al. (1993) compare *Kolopsis sp. with 
*K. yperus, a taxon from Miocene deposits 
disconformably overlying those that contain the 
Alcoota Local Fauna in Northern Territory. They 
contend that *Kolopsis sp. lies morphologically 
between *K. torus (Alcoota) and *K. yperus, thus 
supporting a Miocene age for *Kolopsis sp. on 
stage-in-evolution criteria. 

The ^Zygomaturus gilli holotype P3/ and 
referred maxilla (Stirton 1957) and more recently 
discovered lower molar (Rich 1976), are more 
pristine in preservation and may actually be 
derived from the immediately overlying Black 
Rock Sand. In their evaluation of zygomaturine 
phylogeny, Murray et al. (1993), place *Z. gilli 
above *K. yperus at the base of the genus 
^Zygomaturus further emphasizing its 
phylogenetic separation from *Kolopsis sp. Rich 
et al. (1991) add 'Palorchestes sp.' to the 
Beaumaris faunal list, but this reference seems to 
come from older identifications of the *Z. gilli 
holotype cited in the synonomy provided by 
Stirton (1957). 

The marine invertebrate fauna of the 
transgressive Black Rock Sand was used by 
Singleton (1941) to typify his Cheltenhamian 
Stage. Despite the similarity of its molluscs to 
those of the succeeding Kalimnan stage, he 
assigned the Cheltenhamian to the Upper 
Miocene, largely on the basis of the occurrence of 
teeth of the cetacean Parasqualodon and the 
nautiloid Aturia not known to survive the 
Miocene, but very likely remanie at Beaumaris as 
stated above. Elsewhere in the Port Phillip and 
Gippsland basins a surface of disconformity of 
similar character, developed on Miocene marine 
strata, is overlain by marine deposits of Kalimnan 
age (including the deposits containing the type 
fauna defining the age at Lake Tyers, Gippsland). 
In Gippsland the Jemmys Point Formation also 
rests on the same phosphatic nodule-bearing 
surface of disconformity (Carter 1978). The 
evidence thus seems to indicate that molluscan 
faunas indicative of the Cheltenhamian and 
Kalimnan stages belong to the same transgressive 
interval. Furthermore Darragh (1985) showed that 
molluscan assemblages typical of the 
Cheltenhamian and Kalimnan are superposed 



84 



R. H. TEDFORD 



within the Jemmys Point Formation outcrops at 
Bunga Creek near Lakes Entrance, Victoria. 
Ludbrook (1973: 253-255) gave an historical 
review of this problem which has become 
entwined with efforts to recognize the Miocene- 
Pliocene boundary. 

For the purposes of this review, a Pliocene age 
is assigned to the initiation of the Kalimnan 
transgression. The earliest molluscan faunas 
associated with this transgression are of 
Cheltenhamian composition. They are associated 
with terrestrial vertebrates only at Beaumaris. If 
*Zygomaturus gilli is autochthonous in the Black 
Rock Sand, it represents the oldest Pliocene taxon 
in southeastern Australia. The marine Grange 
Burn Coquina, with an occurrence of the kangaroo 
*Kurrabi sp. (Forsyth's Bank Local Fauna), 
contains a molluscan fauna of Kalimnan type 
(Ludbook 1973) representing a later phase of the 
Pliocene transgression. This unit is not well 
constrained magnetically (Fig. 2 and Whitelaw 
1991). 

Dasyuridae 

Most species are small so that the record of this 
group is biased toward those sites that have been 
screen-washed. Some living genera have Pliocene 
records: Sminthopsis sp. cf. S. mac r our a 
(Fisherman's Cliff), Antechinus sp. (Hamilton, 
Fisherman's Cliff, Dog Rocks), Satanellus 
hallucatus (Fisherman's Cliff), Dasyurus sp. 
(Fisherman's Cliff, Dog Rocks), and Dasyuroides 
*achilpata (Fisherman's Cliff). The living 
Sminthopsis crassicaudata has a medial 
Pleistocene record at Hine's Quarry. The record of 
the largest dasyurid, Sarcophilus, begins in the 
early Pliocene with an unallocated species 
(Parwan) followed by a relatively primitive 
species, S. *moornaensis (Fisherman's Cliff), in 
the late Pliocene. Representatives of the living 
species, or its close ally S. *laniarius (a 
subspecies of S. harrisii to some, Werdelin 1 987), 
occur in early Pleistocene deposits (Limeburner' s 
Point). *Glaucodon ballaratensis, which appears 
to be a member of the Tasmanian Devil clade, 
occurs only in deposits at Smeaton, Victoria, that 
have a Late Pliocene maximum age (Turnbull et 
al. 1993). This genus is thus overlapped in range 
by its sister taxon Sarcophilus. 

Thylacinidae 

Curiously, there is no Pliocene record of 
thylacines in southeastern Australia but the genus 
is recorded in the Miocene Curramulka Local 
Fauna of South Australia (Pledge 1992). There is 



a long record of the genus extending from the 
Miocene into the late Pleistocene in northern 
Australia and New Guinea. The Nelson Bay 
species, close to the modern Thylacinus 
cynocephalus, may be the earliest known 
occurrence of this recently extinct taxon. 

Peramelidae 

This family is also poorly represented in the 
record, very likely because all members are small 
animals. Unidentified peramelids occur at 
Hamilton and Fisherman's Cliff, but at Dog 
Rocks, at the close of the Pliocene, Perameles sp. 
and Isoodon sp. are both recorded. 

Thylacomyidae 

There are no records of bilbies in southeastern 
Australia until the late Pleistocene when Macrotis 
is present in the Murray Basin. 

Thylacoleonidae 

Fragments of the dentition and skeleton of 
*Thylacoleo sp. were recovered with the 
Curramulka Local Fauna (Pledge 1 992) of South 
Australia. The nearby Town Cave produced a 
single P3/, the type of *f. hilli Pledge 1977, which 
is the most primitive member of the genus and 
could be a contemporary of the Curramulka 
assemblage. Unfortunately the fragments from 
these sites do not include comparable elements, 
nevertheless the records do indicate the presence 
of the genus prior to its Pliocene appearance in 
southeastern Australia. This record begins with 
*Thylacoleo sp. at the late Pliocene Bone Gulch 
site in the Murray Basin. Material that can be 
identified with the widespread Pleistocene species, 
*T. carnifex, occurs in the early Pleistocene Duck 
Ponds Local Fauna. 

Vombatidae 

Pledge (1992) records Vombatus, Phascolonus 
and 'Phascolomys" sp., cf. */V medius from the 
later Miocene Curramulka Local Fauna of coastal 
South Australia. Early Pliocene representatives of 
Vombatus are known at Boxlea and Coimadai, 
including V. ^parvus as well as a larger form, V. 
sp. Vombatus ursinus is also present at Dog 
Rocks by the late Pliocene. Lasiorhinus is present 
by medial Pliocene time in the Murray Basin 
(Fisherman's Cliff) and a large species, 
comparable to u Phascolomys' *medius, is present 
in the Port Phillip Basin (Limeburner's Point) by 
the medial Pleistocene. Giant wombats also occur 
in the late Pliocene, *Phascolonus sp. (Dog 



PLIO-PLEISTOCENE MAMMALS OF SOUTHEASTERN AUSTRALIA 



85 



Rocks) and *Ramsayia sp. cf. R. magna (Bone 
Gulch). 

Diprotodontidae 

Remains of these large animals are present at 
nearly all sites providing a more continuous record 
than for other groups except the Macropodidae. 
There is considerable late Cainozoic diversity 
within the family, perhaps denoting a span of 
maximum cladogenesis within the group. The 
diprotodontid record in southeastern Australia in 
the Pliocene includes the diprotodontines 
*Euowenia sp. (Coimadai), a new genus at 
Hamilton, and, perhaps, *Diprotodon itself at 
Fisherman's Cliff and Dog Rocks. The earliest 
record of *Zygomaturus, Z. gilli, occurs in the 
Black Rock Sand at Beaumaris in the Port Phillip 
Basin. This is here regarded as earliest Pliocene as 
discussed above. Other Pliocene occurrences of 
*Zygomaturus sp. are known at Coimadai and 
Dog Rocks. *Zygomaturus trilobus and 
*Diprotodon sp. are present at Nelson Bay in the 
early Pleistocene. By medial Pleistocene time 
*Diprotodon optatum is present at Duck Ponds 
(as 'D. longiceps 1 ), Limeburner's Point and 
Hine's Quarry. The continent-wide diprotodontid 
record shows a reduction in diversity at the close 
of the Pliocene with only *Diprotodon optatum 
and *Zygomaturus trilobus extending into the late 
Pleistocene. 

Palorchestidae 

A relatively primitive early Pliocene species, of 
*Palorchestes, perhaps close to the northern 
Australian Miocene P. painei, occurs at Hamilton. 
A similar form occurs in the later Miocene 
Curramulka Local Fauna of South Australia 
(Pledge 1992). The small *P. parvus is present in 
the early Pleistocene (Nelson Bay) and this taxon 
survives into the late Pleistocene at Strathdownie 
Cave in western Victoria (Gill 1957b). There is no 
early record of the giant *P. azael in southeastern 
Australia although it occurs there in the late 
Pleistocene. 

Phascolarctidae 

Koalas are diverse in Miocene deposits of 
interior Australia, but the earliest representative of 
the genus Phascolarctos occurs in the later 
Miocene Curramulka Local Fauna of coastal 
South Australia (Pledge 1992). A primitive early 
Pliocene species, P. *maris, occurs in the shallow- 
water marine Loxton Sands in the Murray Basin 
(Sunlands Local Fauna, Pledge 1987). A species 



of koala close to the living Phascolarctos cinereus 
is present in the late Pleistocene of southeastern 
Australia. The genus is not recorded in intervening 
deposits. 

Ektopodontidae 

Species of the genus *Darcius are known only 
from Hamilton and another genus of this family 
survived into the early Pleistocene at Nelson Bay 
(M. Whitelaw, pers. comm. 1993) representing a 
relictual occurrence of an important Miocene 
group. 

Phalangeridae 

Members of this family occur in the early 
Pliocene: Trichosurus sp. at Boxlea, and T. 
*hamiltonensis and Strigocuscus *notalis occur 
together at Hamilton. In the late Pliocene, 
Phalanger sp. is recorded at Dog Rocks. 

Petauridae 

Forms close to living small gliders are known 
from Hamilton largely because of the intensive 
screen-washing there. A Petaurus sp., close to P. 
norfolcensis, also occurs in the late Miocene 
Curramulka Local Fauna of South Australia 
(Pledge 1992). Pseudocheirines are somewhat 
larger forms and thus have a wider record before 
the late Pleistocene in southeastern Australia. The 
early Pliocene Hamilton Local Fauna contains two 
extinct Pseudocheirus species, one that appears to 
be related to the living Petauroides 
(Pseudocheirus *stirtoni) and the other to 
members of the living subgenus Pseudocheirus 
(P. *marshalli). Two unidentified species of 
Pseudocheirus are present in the late Pliocene 
(Dog Rocks) and a species close to the living P. 
peregrinus is present in the early Pleistocene at 
Nelson Bay. The latter site also contains a new 
genus of giant pseudocheirine. Additional early 
Pliocene diversity in the ringtail group is also 
indicated by *Pseudokoala erlita, known only 
from Hamilton. 

Burramyidae 

In the well-sampled Hamilton site a species of 
Burramys (B. *triradiatus) occurs in a rainforest 
setting near sea level. This genus is not further 
represented in southeastern Australia until the late 
Pleistocene when the living B. parvus appears in 
superficial deposits in caves in the Buchan district, 
Victoria, at about 600 m elevation. In the 
Holocene it retreated to above 1300 m to be 
associated with alpine environments. 



86 



R. H.TEDFORD 



Potoroidae 

This family has a long Neogene record in 
central and northern Australia, the living genera 
appear in the Miocene in that region. The recently 
described Curramulka Local Fauna (Pledge 1992) 
of later Miocene age also contains an early record 
of Potorous. Hypsiprymnodontines, both 
Hypsiprymnodon sp. and the larger *Propleopus 
sp., occur in the early Pliocene (Hamilton), and 
the latter also at Boxlea, perhaps the earliest 
record of the genus. *Milliyowi bunganditj, a 
newly recognized potoroid of uncertain affinities, 
occurs only at Hamilton indicating some diversity 
within the group in the coastal early Pliocene. 
Bettongia sp. is known in the medial Pliocene in 
the Murray Basin (Fisherman's Cliff). By the 
close of the Pliocene Potorous sp. and a form near 
Bettongia are present in the Port Phillip Basin at 
Dog Rocks. These potoroid genera are common in 
the late Pleistocene of southeastern Australia. 

Macropodidae 

By Pliocene time most of the living genera of 
Macropodinae had appeared in the Australian 
record, as had most of those representing the 
diverse Sthenurinae. There are no living 
macropodine genera in Miocene sites. The record 
in southeastern Australia documents the loss of 
several early genera as well as early species of 
surviving genera during the later Pliocene. Genera 
whose living species have northern Australian and 
New Guinea distributions also fail to persist in the 
southeast beyond the medial Pliocene. 

Sthenurine kangaroos such as *Troposodon and 
*Simosthenurus have later Miocene records at 
Curramulka in South Australia and early Pliocene 
records at Hamilton. *Troposodon did not persist 
in southeastern Australia, but survived into the 
late Pleistocene in northern and eastern Australia. 
*Sthenurus appears by the medial Pliocene 
(Fisherman's Cliff), and continues into the late 
Pliocene (Bone Gulch) and the late Pleistocene in 
the Murray Basin. Species of Sthenurus are 
present in the coastal regions in the late 
Pleistocene although they are not as diverse as in 
the inland. *Simo sthenurus species, such as *S. 
mccoyi, that are close to later Pleistocene forms, 
are present in the medial Pleistocene of the Otway 
Basin. Species of this genus are most numerous in 
the coastal districts of southeastern Australia 
during the late Pleistocene. 

Macropodines of the early Pliocene of 
southeastern Australia include extinct genera that 
did not persist into the later Pliocene; extinct 
genera whose early species did not persist into the 



Pleistocene; living genera whose ranges contracted 
so that they were no longer present in the 
southeastern Australian Pleistocene, and living 
genera that continue to exist in southeastern 
Australia. Examples of the first are species of 
*Kurrabi (*K. pelchenorum) at Hamilton and 
*Kurrabi sp. at Coimadai and Forsyth's Bank 
(Flannery et al. 1992). The second case is 
exemplified by early *Protemnodon species such 
as the *P. chinchillaensis that occurs in Kalimnan 
strata at Lake Tyers (originally identified as *P. 
otibandus, Plane 1972, reidentified by Rich et al. 
1991) and *P. cf otibandus at Hamilton. The last 
recorded occurrence in southeastern Australia of 
such early species of *Protemnodon, is *P. devisi 
from the Fisherman's Cliff Local Fauna of late 
Pliocene age in the Murray Basin. 

Dorcopsis (D. * winter cookoruni) and cf. 
Dendrolagus occur in the early Pliocene of the 
Otway Basin (Hamilton). Dorcopsis is also 
present in the western Murray Basin (Sunlands) in 
the early Pliocene. These living genera do not have 
later records in southeastern Australia. On the 
other hand species of Thylogale (T. *ignis), 
perhaps Wallabia and Macropus (Notamacropus) 
are recorded in the early Pliocene of the Otway 
Basin (Hamilton). In medial Pliocene time 
Lagostrophus sp. cf. L. fasciatus, Petrogale sp. 
and Macropus (Osphranter) sp. are present in the 
Murray Basin (Fisherman's Cliff). 

Species of Macropus (Macropus) may be 
present in the early Pliocene (Coimadai), but a 
more secure record is present only late in the 
epoch at Dog Rocks in the Port Phillip Basin 
where species close to the living grey kangaroos 
(both M. giganteus and M . fuliginosus) occur. The 
occurrence of a skeleton of Macropus giganteus 
beneath basalt in the Great Buninyong Estate 
Mine, Victoria, implies significant antiquity for 
this taxon, perhaps predating 2.5 Ma, the age of 
basalt flows in the vicinity (Rich et al. 1991). The 
Dog Rocks site also produced the earliest record 
of Wallabia bicolor and a wallaby near M. 
(Notamacropus) irma, presently confined to 
Western Australia. A large *Protemnodon, 
comparable to *P. analc, succeeds earlier species 
in the late Pliocene (Dog Rocks). 

By the beginning of the Pleistocene Macropus 
(M.) *titan appears with * Protemnodon sp. cf. P. 
analc at Duck Ponds, the earliest record of 
coexistence of two of the more common 
Pleistocene macropodid species. At Nelson's Bay 
the occurrence of *Baringa nelsonensis indicates 
survival into the Pleistocene of a macropodine also 
known in the late Miocene Curramulka Local 
Fauna of South Australia. 



PLIO-PLEISTOCENE MAMMALS OF SOUTHEASTERN AUSTRALIA 



87 



The small samples available of medial 
Pleistocene faunas (Limeburner's Point and Hine's 
Quarry) indicate the continued occurrence of 
larger M. (Macropus) and *Protemnodon, the 
presence of smaller Macropus species, Wallabia 
sp. cf. W. bicolor and Macropus (Notamacropus) 
sp. cf. M. (N.) parryi at Limeburner's Point and 
the possible survival of the northern Australian 
Pliocene *Prionotemnus sp. at Hine's Quarry. 
Lack of Murray Basin sites of earlier Pleistocene 
age prevent establishment of the ranges of 
Lagorchestes and Onychogalea, genera that were 
common there in the late Pleistocene and 
Holocene. 

Muridae 

The array of small mammals obtained from the 
Hamilton site through intensive screen-washing 
lends credence to the absence of murid rodents 
there. Likewise the Miocene Curramulka Local 
Fauna lacks rodents although it has rodent-sized 
small marsupials. They have been reported from 
Parwan whose estimated age is 4.0 Ma., but 
supporting collections cannot be found in the 
Museum of Victoria. Thus the earliest verfied 
occurrence in southeastern Australia is the 
material from the medial Pliocene Fisherman's 
Cliff Local Fauna in the Murray Basin. At that 
time the murids had achieved considerable 
diversity (Crabb 1976) and living genera such as 
Pseudomys, Leggadina, Leporillus and Notomys 
have been identified there. Pseudomys is also 
present in the late Pliocene (Dog Rocks) of the 
Port Phillip Basin. 



Implications of the faunal record 

Despite its obvious imperfections, the Pliocene- 
Pleistocene faunal sequence in the Murray and 
coastal basins of Victoria constitutes the most 
complete record of faunal change for that interval 
presently known on the continent. Central 
Australia (Lake Eyre Basin) and northeastern 
Australia (Charters Towers area) contain 
important dated early Pliocene assemblages, but 
large gaps separate these from younger Pleistocene 
faunas in the same districts. The record from the 
Murray Basin has a similar gap, but its medial 
Pliocene record nicely complements the coastal 
sequence. This is important as faunas on either 
side of this gap differ significantly through 
extinction, evolution and zoogeographic changes 
of their components. The evidence from the 



Murray and coastal basins not only supports this 
conclusion, but allows us to chart the changes and 
provides a rough chronology for them. 

Reassembling the faunal records temporally, 
three responses seem to be in play during a span 
of faunal change: 1) extinction of pre-existing 
taxa, 2) survival and, in some cases, evolution of 
pre-existing taxa, and 3) changes in geographic 
range of surviving taxa so that they become locally 
extinct. As the range chart (Fig. 3) indicates, 
important faunal turn-over occurred during the 
medial to late Pliocene. In the southeastern 
Australian early Pliocene a number of living and 
Pleistocene genera are already present, including 
Antechinus, Sarcophilus, Strigocuscus, 
Vombatus, *Diprotodon, Hypsiprymnodon, 
*Propleopus, Dendrolagus, Dorcopsis, 
*Protemnodon, *Troposodon, *Simosthenurus, 
Thylogale, Wallabia and Macropus 
(Notamacropus). *Euowenia, *Darcius, 
*Troposodon and *Kurrabi have records limited 
to the early Pliocene in southeastern Australia, 
while Strigocuscus, Hypsiprymnodon, Dorcopsis 
and Dendrolagus are not known in the region 
after the early Pliocene. However species of these 
living genera are restricted to northern Australia 
and New Guinea today, and *Tropososon 
continued to be represented in eastern and 
northern Australian faunas into the late 
Pleistocene. 

During the medial Pliocene in the Murray Basin 
additional living genera occur, some represented 
by species that appear limited to the Pliocene: 
Sminthopsis, Dasyuroides and Dasyurus, 
Satanellus, Lasiorhinus, Bettongia, Macropus 
(Osphranter), Petrogale and Lagostrophus. 
Dasyuroides and Satanellus are not known in 
younger faunas, but occur in northern Australia 
today. *Sthenurus occurs first in the Murray Basin 
and becomes a member of the coastal assemblage 
by the late Pleistocene. 

In the late Pliocene some living species, or 
forms close to them, appear in the Port Phillip 
Basin: Vombatus ursinus, Wallabia sp. cf. W. 
bicolor, Macropus (Macropus) sp. cf. M. (M.) 
giganteus, M. (M.) sp. cf. M. (M.)fuliginosus and 
M. (Notamacropus) sp. cf. M (N.) irma. Some 
extinct genera must have undergone species level 
changes during this interval for by the beginning 
of the Pleistocene *Diprotodon optatum, 
*Zygomaturus trilobus, *Thylacoleo carnifex, and 
*Protemnodon anak appear in coastal Victoria. 
*Ramsayia magna also appears in the late 
Pliocene of the Murray Basin. These taxa, along 
with Macropus (Macropus) *titan, are among the 



88 



R. H.TEDFORD 



most widespread components of the Pleistocene 
faunas of southeastern Australia. 

In early to medial Pleistocene time some 
Tertiary relicts have their last appearances: 
*Ektopodontidae, *Baringa and perhaps 
*Prionotemnus. Further diversity is evident in 
living genera [e.g. Macropus (Notamacropus) sp. 
cf. M. (/V.) parry i] and extinct forms such as 
"Phascolomys" cf. *medius are present. Extinct 
genera such as *Simosthenurus show initial 
phases of speciation (*£. mccoyi) heralding their 
later Pleistocene radiation. 



Palaeoecological interpretations from the 
mammalian fauna 

Despite its inadequacies, the faunal evidence 
from southeastern Australia supports three general 
ecological conclusions. The first is that early 
Pliocene sites have genera whose living species 
are now members of the tropical rainforest 
assemblage, notably Hypsiprymnodon, 
Dendrolagus, Dorcopsis and Strigocuscus from 
Hamilton and Dorcopsis at Sunlands, suggesting 
similar environments prevailed in the Otway and 
western Murray basins at that time. Such taxa do 
not appear in younger Pliocene sediments of the 
coastal region, although there is a clear signature 
of relatively humid later Pliocene environments 
with Vombatus, Potorous, Wallabia, Macropus 
(Notamacropus), Thylogale and arboreal 
phalangerids and petaurids, all taxa that today 
inhabit wet sclerophyll forest. 

Secondly there is an ecological contrast between 
the inland Murray Basin and coastal Victoria 
when comparable records are available in the 
medial to late Pliocene. Arboreal taxa are more 
commonly recorded on the coast, less frequently 
inland, judging from comparably collected sites 
(compare Fisherman's Cliff and Dog Rocks). 
Certain genera whose living species were widely 
distributed in the modern arid zone in immediately 
pre-European times (Dasyuroides, Lasiorhinus, 
Leggadina, Leporillus), or are strongly 
represented in the arid-zone today (Macropus 
(Osphranter), Petrogale, Notomys, Pseudomys), 
are found in the Murray Basin in the medial to 
late Pliocene suggesting a climatic gradient 
similar in orientation to that of today. 

A third conclusion is that the Pliocene 
assemblages of both the coastal and inland sites 
contain associations of living genera not known 
today. Flannery et at. (1992) point out the 
association of Trichosurus and Strigocuscus at 



Hamilton as an example of taxa whose ranges do 
not overlap today, and the coexistence of 
Hypsiprymnodon, Dorcopsis and Burramys is 
another example from the same assemblage. 
Likewise, in the medial Pliocene of the Murray 
Basin (Fisherman's Cliff), Lagostrophus, 
Dasyuroides, and Satanellus are genera now 
widely separated zoogeographically. Such 
coexistence of taxa in communities that lack 
modern analogues (the "disharmonious 
associations" of Lundelius 1983) imply unusually 
patterned environments or subsequent adaptations 
of species of these genera to habitats occupied by 
their living representatives. The latter is clearly 
the case for the species of Burramys, as already 
mentioned. 

Dated early Pliocene assemblages are known in 
central Australia (Tirari Formation local faunas, 
Lake Eyre Basin, Tedford and Wells 1992) and 
northeastern Australia (Bluff Downs Local Fauna, 
Queensland, Archer and Wade 1976). Both 
contain rodents and on this basis may slightly 
post-date the Hamilton Local Fauna. They are 
constrained by local geochronological indicators 
that suggest an age near 4 Ma for both areas. 
Except for the presence of a large Dendrolagus 
and the phascolarctid *Koobor these northern 
faunas are dominated by terrestrial forms. The 
Pliocene genera *Euowenia and *Kurrabi are 
shared with southeast Australia and *Diprotodon 
also occurs in the Tirari faunas. There are early 
Pliocene occurrences of living genera such as 
Lagorchestes, Macropus (Osphranter) and 
Macropus (Macropus), and extinct genera 
*Thylacoleo, *Phascolonus, and *Sthenurus in 
the northern assemblages, the latter predates its 
record in southeastern Australia. The association 
of Dendrolagus with Macropus species and 
Lagorchestes has no modern equivalent but could 
indicate a vegetational mosaic, possibly riparian 
gallery forest and more open associations on river 
valley interfluves, with the remains of taxa from 
parts of the mosiac being brought together by 
fluviatile agencies. The comparison between 
southeastern and northern Australian Pliocene 
faunas suggests different zoogeographic 
patterning, broader mesic environments and less 
climatic diversity than present today. 



Palaeoecological interpretations from geological 
history and the palaeobotanical record 

Comparison of the faunal succession and its 
palaeoecological interpretation with physical and 



PLIO-PLEISTOCENE MAMMALS OF SOUTHEASTERN AUSTRALIA 



89 



other biological events in southeastern Australia 
indicates sufficient congruence in the timing and 
nature of change to suggest global controls of the 
events. Brown and Stephenson's (1991) masterful 
synthesis of the history of the Murray Basin is 
heavily drawn upon in the following account. 

A hiatus of about 4 M.y. accompanies the late 
Neogene regression in the Murray and coastal 
basins during which these basins were emergent 
for much of the late Miocene. The sea withdrew 
beyond the present coastline, possibly to the 
continental edge, and the Murray Basin 
environment in particular became continental. 
Martin's (1987) study of the vegetational changes 
in the Lachlan Valley in the eastern Murray Basin 
shows that this period of emergence is 
accompanied by the first appearance of wet 
sclerophyll forest following a long Palaeogene and 
early Neogene occurrence of rainforest formations 
in the basin. The Curramulka Local Fauna of 
Yorke Penninsula, South Australia belongs to this 
regression. The cave system from which the fauna 
was recovered was open to the surface and 
collecting terrestrial debris during a period of 
lower water table. Lack of rainforest taxa in the 
assemblage perhaps reflects the drier environment 
in the karstic uplands of the peninsula. 

The subsequent Pliocene transgression in the 
Murray Basin was accompanied by deposition of 
the shallow-marine Bookpurnong Beds followed 
by the marine Loxton Sand. During this high stand 
of sea level, rainforest vegetation redeveloped in 
the eastern part of the Basin. Martin (1937) has 
shown that although this forest contains 
Nothofagus species, only the temperate species 
groups are represented and these may have been 
restricted to the uplands surrounding the basin and 
to the more protected river valleys. Judged by its 
mammalian inhabitants, early Pliocene lowland 
rainforest of southeastern Australia may have 
resembled present-day rainforest of northeastern 
Australia. The few terrestrial fossils obtained from 
the Loxton Sand in the western part of the basin 
(Sunlands Local Fauna, Pledge 1985, 1987) 
include a large koala {Phascolarctos *maris), 
Dorcopsis sp. and a *zygomaturine diprotodontid. 
This assemblage may be approximately correlative 
with the Hamilton and Forsyth's Bank local 
faunas in the Otway Basin and the Lake Tyers 
Local Fauna in the Gippsland Basin, all occur in 
the transgressive phase of sedimentation (the 
Kalimnan Stage) in their respective basins. These 
assemblages are the only Pliocene faunas 
containing rainforest taxa. 

When the sea withdrew from the Murray Basin 



in the early Pliocene, strandline dunes followed 
(Parilla Sand) and finally fluviatile deposits 
(Moorna Sand) penetrated interdune regions as 
regional drainage was re-established. Increasingly 
continental conditions in the Murray Basin were 
accompanied by the final demise of the rainforest 
and myrtaceous sclerophyll vegetation reappeared 
in the basin (Martin 1987). The Fisherman's Cliff 
Local Fauna was contemporaneous with this phase 
of regression and reflects the change to drier 
environments. Judged by the development of 
silicified ferruginous soils on the stabilized 
regression surface (the Karoonda Surface of 
Firman 1966), the climate included higher, and 
perhaps less seasonal, rainfall and temperatures 
higher than present. 

In the Otway Basin the regression was 
accentuated by tectonic and thermal uplift 
accompanying the onset of the volcanism that built 
the Newer Volcanic plateau of western Victoria. 
The Port Phillip Basin forms part of the hinge 
zone between the Otway and Gippsland Basins. 
Its geological history resembles the latter, but late 
Neogene sequences are thinner and Newer 
Volcanic flows punctuated the section during the 
Pliocene. In the Port Phillip Basin the early 
Pliocene regression was accompanied by 
dissection of the Kalimnan transgressive sands 
(Moorabool Viaduct Sand). Backfilling of the 
erosion surface by thin marine deposits containing 
faunas attributed to the Werrikooian Stage (Carter 
1985) and terrestrial deposits with the Dog Rocks 
Local Fauna, represent a short-lived late Pliocene 
transgression. This has its equivalent in the 
Murray Basin in the Norwest Bend Formation that 
was limited to the westernmost part of the basin. 
This was the last marine incursion onto the 
southern part of the continent. Thereafter 
Quaternary eustatic transgressions were confined 
to more coastal districts. 

In the Murray Basin late Pliocene uplift of the 
coastal Pinnaroo block provided a tectonic dam 
limiting the sea and blocking the fluviatile 
drainage of the Basin. These movements 
impounded the lower drainage of the basin's trunk 
stream (the Murray) and lacustrine claystone 
(Blanchetown Clay) accumulated in this large lake 
during late Pliocene through the early Pleistocene 
time (essentially the duration of the Matuyama 
Chron, An et al. 1986). Wet sclerophyll vegetation 
again occupied the eastern Murray Basin, 
responding to the humid conditions engendered by 
the large lake, itself mimicking the climatic effects 
of a marine transgression. Fluviatile sand bodies 
reached into the lacustrine deposits as the lake 



90 



R. H.TEDFORD 



level rose and fell in response to climatic 
perturbations. One of these fluviatile units 
(collectively the Chowilla Sand) contains the Bone 
Gulch Local Fauna of late Pliocene age. By the 
early Pleistocene rainfall diminished, evaporation 
rates increased and dolomitic limestones 
(Bungunnia Limestone) were deposited in the 
western part of the lake basin before an outlet was 
finally formed and the lake drained at about 0.7 
Ma. The resulting increase in gradient entrenched 
the Murray River in its present course. Falling 
groundwater levels provided large expanses of 
unvegetated terrain that were available to aeolian 
processes under the increasingly drier and more 
seasonal climates of the Quaternary. As forest 
cover dwindled in the Pleistocene, woodland and 
grassland/herb fields developed as precipitation 
fell to modern values and beyond (Martin 1987). 
A fossil vertebrate record of this span of 
environmental change has not yet been found in 
the Murray Basin, the record continues there in 
the late Pleistocene after a substantial hiatus. 

In the coastal district of Victoria late Pliocene 
through medial Pleistocene mammal faunas are 
present, and although the evidence is still sparse, 
it gives insight into the faunal changes during a 
span of important environmental change beyond 
that recorded in the Murray Basin. The 
Limeburner's Point and Hine's Quarry local 
faunas show that taxa common to the Late 
Pleistocene faunas of southeastern Australia 
already formed communities in the medial 
Pleistocene and continued through the rest of the 
Pleistocene without major change. Only the 
sthenurines seem to show increasing diversity of 
form as though this 0.7 M.y. span was a period of 
intense speciation. 

Comparative palaeoenvironmental evidence is 
also available from the circum-Antarctic ocean 
(Hodell and Venz 1992 and references therein) and 
the Antarctic continent itself (Ishman and Rieck 
1992 and references therein) covering the Pliocene 
and early to medial Pleistocene. These data 
indicate that the early to medial Pliocene seas were 
relatively warm with high sea-levels drowning 
coastal valleys in Antarctica and corresponding to 
the transgressive phase of Pliocene history in 
southeastern Australia. At approximately 2.6-2.4 
Ma, at the beginning of the late Pliocene, a 
significant shift in 5 IX of both planktic and 
benthonic foraminifers toward positive values 
heralds the beginning of a major cooling in 
Antarctica, an event coincident with the initiation 
of the first major build-up of ice at high latitudes 
in the northern hemisphere. The resulting sealevel 



drop is recorded as the post-Kalimnan regression 
in southeastern Australia. Cooler climates marked 
the demise of significant tropical rainforest in 
southeastern Australia and the return to temperate 
sclerophyll vegetation on the coast and more open 
formations inland. This early glacial-interglacial 
cycling was of irregular frequency allowing minor 
transgression (Werrikooian) in southeastern 
Australia, but by 1 .4 Ma, in the early Pleistocene, 
a strong 0.041 M.y. cycle becomes evident in the 
5 I8 signal and this feature, associated with 
variation in the earth's obliquity cycle, persists 
throughout the Pleistocene. Thus early in the 
epoch Australia's biota was subjected to the 
climatic oscillations that were to typify the 
Pleistocene. Direct evidence of these cycles from 
the continent is woefully incomplete, but 
significant mountain glaciation in Tasmania 
reaches into the Matuyama Chron (Colhoun and 
Fitzsimons 1990) in agreement with the limits of 
such effects as determined from Antarctica. 



Conclusions 

Improved chronological resolution of the 
Pliocene and Pleistocene mammal-bearing 
deposits of southeastern Australia permits 
determination of the temporal succession of taxa 
and assessment of the zoogeographic and 
ecological significance of their occurrence. More 
comprehensive palaeomagnetic data, calibrated 
mostly by reference to radioisotopically dated 
flows of the Newer Volcanic field in Victoria, and 
more limited data from the Murray Basin, provide 
a chronology for the faunal succession from early 
Pliocene into medial Pleistocene time. Nowhere 
else in Australia is such a detailed representation 
of this interval to be found. Despite the 
preservational, taphonomic and palaeoecological 
biases of the record, it provides the best 
documented model of mammal faunal change at 
the end of the Australian Cainozoic. 

Early Pliocene faunas in both the Murray and 
Otway Basins have greater affinity with 
Pleistocene and Holocene faunas than they do with 
known Miocene assemblages. Younger strata 
within the Pliocene transgression show many 
genera that represent the earliest members of 
lineages that persist through the Pleistocene and 
into the Holocene. In that sense the 'modern' 
fauna appears in the fossil record at the beginning 
of the Pliocene, but for some genera, their 
evolutionary appearance may considerably predate 
this first local appearance as suggested by the 



PLIO-PLEISTOCENE MAMMALS OF SOUTHEASTERN AUSTRALIA 



91 



Curramulka Local Fauna (Pledge 1 992) of nearby 
South Australia. The late Miocene hiatus in the 
southeastern Australian basins limits opportunities 
to discover the actual historical record there. 

Early Pliocene faunas of southeastern Australia 
contain several genera that appear not to be 
members of younger assemblages including the 
diprotodontid *Euowenia, the potoroid *Milliyowi, 
the macropodine *Kurrabi and perhaps the 
ektopodontid *Darcius. During the span of the 
early Pliocene the genera *Zygomaturus, 
*Diprotodon, and *Propleopus, appear in 
southeastern Australia along with species, some 
extinct, of the living Antechinus, Sarcophilus, 
Vombatus, Trichosurus, Thylogale, Macropus 
(Notamacropus), Wallabia, Hypsiprymnodon, 
Dorcopsis, Dendrolagus, and Strigocuscus. The 
latter four rainforest genera are unrecorded in later 
deposits in southeastern Australia, and the co- 
occurrence of species of some of these living 
genera represent associations unknown in living 
faunas. The early Pliocene marine transgression in 
the Murray Basin and coastal basins of Victoria 
brought humid conditions onto the continent and 
rainforest was present in southeastern Australia. 

Medial Pliocene regression restored continental 
environments to the Murray Basin and species of 
the living Dasyuroides, Satanellus, Lasiorhinus, 
Bettongia, Macropus (Osphranter), Petrogale, 
Lagostrophus, Notomys, Leggadina and 
Leporillus appear in response to drier 
environments. Establishment of a large lake 
system in the western and central part of the 
Murray Basin in the medial Pliocene allowed wet 
sclerophyll vegetation to invade the basin. These 
conditions persisted into the early Pleistocene 
before the lake system was drained and drier 
shrubland vegetation became more widespread. 

The late Pliocene and early Pleistocene appears 
to have been a time when living species, or forms 
close to them, appear in the coastal regions. 
Species such as Vombatus ursinus, Wallabia sp. 
cf. W. bicolor, Macropus (Macropus) giganteus, 



M.(M) fidiginosus and M. (Notamacropus) irma, 
or closely allied taxa, appear in the late Pliocene. 
By the beginning of the Pleistocene a number of 
extinct species appear in the coastal region, 
Sarcophilus *laniarius, *Diprotodon optatum, 
*Zygomaturus trilobus, *Thylacoleo carnifex, 
*Protemnodon anak and Macropus *titan, all of 
which persist through the Pleistocene as the 
characteristic large mammal suite in most 
Quaternary assemblages. 

The 'modernization' of the mammal fauna 
during the late Pliocene to early Pleistocene 
interval corresponds with the increasing 
continentality of southeastern Australia and to the 
climatic changes toward modern environments 
that are observed world-wide as the rate of glacial- 
age climatic cycling begins to increase. The faunal 
turnover at the close of the Pliocene, roughly 
coincident with the initiation of bipolar glaciation, 
also introduced taxa into the region that had no 
close relatives there and that seem to be 
immigrants from drier environments already 
established elsewhere. Earlier Pliocene histories 
for some of these [e.g. Lagorchestes, Macropus 
(Macropus) and Macropus (Osphranter)] are 
present in inland Australia. Later in the Pliocene 
and Pleistocene environmental contrasts between 
coastal and interior southeastern Australia yield 
zoogeographic patterns similar to those of the 
present-day. 



Acknowledgments 

This synthesis would have been impossible to advance 
without the kind interaction with Dr. Mick Whitelaw 
whose work provided the crucial data. His generosity in 
keeping me informed of his results in advance of 
publication allowed timely preparation of this work. 
Reviewers also materially helped the presentation, 
especially Dr. Alex Baynes, whose careful critique 
helped sharpen the paper and remove inaccuracies in the 
original work. 



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STUDIES OF THE LATE CAINZOZOIC DIPROTODONTID MARSUPIALS 
OF AUSTRALIA. 4. THE BACCHUS MARSH DIPROTODONS - GEOLOGY, 

SEDIMENTOLOGY AND TAPHONOMY 

John Long & Brian Mackness 



Summary 

The remains of at least twenty individuals of Diprotodon were excavated from the Hine's Quarry 
Fossil Site near Bacchus Marsh, Victoria over the 1973 and 1979 field seasons. Sediments 
entombing the fossils consist of closed framework sands and grits with interspersed clay lenses, 
which sit on top of the Tertiary Werribee Foundation. Sedimentary structures and comparison with 
recent sediments of the area indicate that the depositional environment was the proximal part of a 
low flow regime, ephemeral run-off system, containing an intercalated erosional screes suggestive 
of times of non-hydraulic influence. Bone orientations are related to channel morphology with 
minimal hydraulic action responsible for poor sorting into Voorhies groups. Bone surface textures 
and fracture patterns indicate possible carnivore gnawing and prolonged subaerial exposure in an 
arid climate. Articulated skeletons are common in the proximal area of the bone bed with reworked 
skeletal material distally. Regional lithologies were reviewed with respect to their contribution to 
the bonebed. 



STUDIES OF THE LATE CAINOZOIC DIPROTODONTID MARSUPIALS OF 

AUSTRALIA. 4. THE BACCHUS MARSH DIPROTODONS — GEOLOGY, 

SEDIMENTOLOGY AND TAPHONOMY. 



JOHN LONG & BRIAN MACKNESS 



LONG, J. & MACKNESS, B. 1994. Studies of the late Cainozoic diprotodontid marsupials of 
Australia. 4. The Bacchus Marsh Diprotodons - Geology, sedimentology and taphonomy. Ret. 
S.Aust. Mus. 27(2): 95-1 10. 

The remains of at least twenty individuals of Diprotodon were excavated from the Hine's 
Quarry Fossil Site near Bacchus Marsh, Victoria over the 1973 and 1979 field seasons. 
Sediments entombing the fossils consist of closed framework sands and grits with interspersed 
clay lenses, which sit on top of the Tertiary Werribee Formation. Sedimentary structures and 
comparison with recent sediments of the area indicate that the depositional environment was the 
proximal part of a low How regime, ephemeral run-off system, containing intercalated erosional 
screes suggestive of times of non-hydraulic influence. Bone orientations are related to channel 
morphology with minimal hydraulic action responsible for poor sorting into Voorhies groups. 
Bone surface textures and fracture patterns indicate possible carnivore gnawing and prolonged 
subaerial exposure in an arid climate. Articulated skeletons are common in the proximal area of 
the bone bed with reworked skeletal material distally. Regional lithologies were reviewed with 
respect to their contribution to the bonebed. 

John Long, Western Australian Museum, Francis Street, Perth, Western Australia. Brian 
Mackness, School of Biological Sciences, University of New South Wales, P.O. Box 1, 
Kensington, NSW 2033. Manuscript received 7 September 1993. 



The remains of diprotodontid marsupials are 
commonly found in the Late Cainozoic deposits of 
Australia but normally comprise isolated elements 
such as jaws, limb bones or teeth. Whole 
skeletons including skull and mandibles are a 
much rarer occurrence but a number have been 
recovered including those of Diprotodon from 
Lake Callabonna (Tedford 1973; Pledge 1993; 
Tedford 1993); Zygomaturus from Mowbray 
Swamp (Scott 1915); Euryzygoma from the Bluff 
Downs Local Fauna (Mackness unpublished 
information) and Neohelos from Bullock Creek 
(Tom Rich pers. comra.). 

A number of Diprotodon bones were first 
discovered at Hine's Quarry, 9 km south-west of 
Bacchus Marsh, central Victoria, in 1973 by Miss 
Kerry Hine. Subsequently the Museum of Victoria 
(then the National Museum of Victoria) began 
excavating a rich concentration of largely 
articulated Diprotodon skeletons, lead by Thomas 
Darragh and K. Simpson (Monash University). In 
1979, further bones were uncovered by quarry 
work. Periodic field work was resumed under Dr. 
Tom Rich, Museum of Victoria up until December 
1979 when the bone bed was completely dug out. 
During this time bone elevations and orientation 
relative to two fixed datum points were measured, 
and sediment samples were taken from the bone 



bed and its surrounding outcrops, for detailed 
analysis. 

Rich (1976) regarded the Hine's Quarry site as 
probably a channel deposit, and from comparison 
with the present level of the Parwan Valley 
suggested that the site could be relatively old. The 
only age control on the site is the Newer Basalt 
which is present within the fossil bearing 
sediments as cobblestones. The Newer Basalt lies 
above other lava flows in the region. The oldest of 
these flows has a maximum date of 4.03 million 
years (Rahman & McDougall 1972), thus giving a 
maximum age range for the deposit as somewhere 
from Pliocene to Recent. 

This paper reports on the taphonomy of this 
fossil site, examining broadly the geological and 
biological factors which indicate the possible 
cause of mortality and subsequent prediagenetic 
events leading up to the death assemblage 
uncovered in the quarry. It is based largely on the 
work of Long (1979, Monash University 3rd Year 
field project). Future papers by Mackness will 
discuss the taxonomy and tooth variation of the 
Bacchus Marsh Diprotodon site as well as its 
associated microfauna. Few taphonomic or 
palaeoecological studies of Australian Pleistocene 
mammal sites have so far been carried out (Horton 
1976; Horton & Samuel 1978; van Huet 1993). 



96 



J. LONG & B. MACKNESS 



Methods 

Bone orientations were measured by compass 
bearing to the long axis on each bone. Elevation 
was measured using a plastic tube filled with a 
dark liquid. The end of this tube was tied to the 
datum pipe, with the level marked and then a 
measuring pole placed on the bone. The free end 
of the tube was moved up or down until the 
marker level on the datum pipe was achieved, and 
then height of fluid level was measured above the 
bone and recorded. 

Field mapping was carried out in the immediate 
vicinity of Hine's Quarry, and sediment samples 
were collected from nearby ephemeral streams and 
eroding gullies for comparison. Sedimentological 
studies were carried out by standard dry sieving 
procedures, thin sectioning, and differential 
thermal analysis (DTA), the latter carried out at 
the Mineral Physics Laboratories of the CSIRO, 
Port Melbourne. Bone surface textures were 



studied with the aid of a scanning electron 
microscope, and analysis of the composition of 
fossil and fresh marsupial bone was determined 
by use of an electron microprobe. 
Sedimentological methods follow those of Folk 
(1961). The orientations of bones collected during 
the 1973 field season were determined from a 
series of detailed photographs taken by Ian 
Stewart. Abbreviations for specimen/sample 
numbers: P, Museum of Victoria; #, Field 
Number; MG, Monash University Department of 
Earth Sciences. 



Geology 

Hine's Quarry is situated in the south-eastern 
corner of the sunken Ballan Graben, which forms 
part of the uplifted block on the west of the 
Rowsley Fault, bounded to the south by the 
Brisbane Ranges, and to the north by the 




FIGURE 1. Local geology of the Hine's Quarry region, 9 km south-west of Bacchus Marsh, central Victoria, 
Australia. 



BACCHUS MARSH DIPROTODON 



97 




i:M 



t 







, * 



m 



M ,^:# : -' 



<*'% 



■' ' ' mf'W 




FIGURE 2. View of Hine's Quarry fossil excavation (arrow points to main bone horizon), looking east from the 
south-western corner of the quarry. 



Lerderderg Ranges (Fenner 1918; Summers 
1923). Parwan Creek and Werribee River cut into 
the soft clays and sandy clays of the Werribee 
Formation (Thomas & Baragwanath 1950; then 
called the Yaloak Formation), draining to the 
eastern lowland of the Werribee Plains. The fossil 
site (Fig. 1) is situated close to the Rowsley Fault 
which is post-Pliocene in age based on the 
evidence of faulted Newer Basalt dated at 4.03 
million years (Rahman & McDougall 1972). 

The fossils occur in the south-eastern corner of 
Hine's Quarry (Fig. 2) in a thin surficial layer of 
reworked sediments derived primarily from the 
underlying Werribee Formation. The Werribee 
Formation is made up of kaolin-rich sediments 
which (Fig. 3) are distinguished by their low 
quartz content, absence of basalt inclusions, and 
more compact texture. Inclusions within the 
Werribee Fm are infrequent, but quartz pebbles 
and gravel lenses are present, derived from nearby 
Permian glacials and uplifted Ordovician flyschoid 
sediments. Two important units of the Werribee 
Formation are exposed near the fossil site, both 
which have contributed to the composition of the 



fossiliferous sediments. The age of the Werribee 
Formation is tentatively placed as being Paleocene 
to Mid Miocene based upon microflora from the 
Lal-Lal coal deposit (Cookson 1954, 1957), and 
relationships with the older basalts (Wellman 
1974). Exposed portions of the Werribee 
Formation at Hine's Quarry are not fossiliferous. 

Ferruginized grits are exposed to the south of 
Hine's Quarry, forming 'Hine's Hill'. These are 
closed framework quartzose, coarse sandstones 
and gravels, representative of fluviatile channel 
sediments with interspersed minor conglomerates 
and cross-bedded sandstones. The unit exposed at 
Hine's Hill is approximately ten metres thick. The 
top of this unit is capped by recent, relatively 
unconsolidated fluviatile conglomerates. The 
ferruginized grits sit conformably upon the fine 
kaolinites exposed in the quarry. No basalt caps 
Hine's Hill, it being of higher elevation than the 
surrounding basalt plain. 

At Werner's Quarry, one kilometre east of 
Hine's Quarry, the ferruginized grits are seen 
above the kaolin and below the Newer Basalt 
flows, confirming that this unit is included within 



98 



J, LONG & B. MACKNESS 




FIGURE 3. Contact between reworked fossiliferous surface sediments and Tertiary Werribee Formation kaolin 
(arrow indicates unconformity). Note coarse basal gravels containing large basalt cobblestones. 



the Werribee Formation. There are palaeosols and 
laterile with accretionary structures, roughly 
spheroidal and inwardly zoned, between the grits 
and basalt. This resembles the Timboon terrain in 
the Parwan valley (Gill 1964). 



Sedimentology 

Although fossil bones were found at two sites 
within Hine's Quarry, the main concentration 
came from the south-eastern corner, the subject of 
this study. Another locality, 'Ian's Site', has 
produced a few bones from the top of the south- 
west face of the quarry. Both sites involve similar 
lithology and arc probably contemporaneous. 

The sediments entombing the main bone 
concentration comprise discontinuous layers of 
clay-rich closed framework sands, gravels and 
silts (Fig. 4). They represent a mixture of both 
depositional and diagenetic events. Large basalt 
cobblestones occur randomly throughout the 
deposit, either as isolated stones without 
associated gravel, or within small conglomeratic 
lenses. 



Below the bone-bearing sediments is the 
diseonformable contact with the kaolinitic clays of 
the Werribee Formation, and above the unit arc 
recent river terrace conglomerates, so close to the 
subsurface as to be incorporated as regolith. The 
fossiliferous unit is up to 3.3 metres thick at the 
distal end of the bone concentration. The 
concentration of Diprotodon remains form the 
only marker horizon within these sediments, 
although spasmodic isolated bones (generally 
macropodid remains) randomly occur at all 
stratigraphic levels. 

Several profiles through the unit were measured 
and described. The discontinuity of sediment 
layers can be clearly seen from these sections (Fig. 
4). Sedimentary structures were generally absent 
apart from horizontal discontinuous stratification, 
lensoidal and festoon cross-stratification. Rarer 
instances of small scale scour and fill channels, 
graded laminar bedding, and steep erosional 
surfaces were encountered. The high clay content 
of the coarser beds is due to secondary seepage of 
clay through the open pores of the sand. These 
sediments arc strongly bimodal, and have been 
analysed with the diagcnctically acquired fine 
fraction omitted. 



BACCHUS MARSH DIPROTODON 



99 



N ■* 



o-t 




sand 



basalt cobble- 



G 


H \ 


r^T' 






• o B o° 






m 


^ 


8p 


- r_ - - 




^r 


W& 




.;"-■-.?-; 


'o°^. 




()»■■■ 


>6 0°Op 




II 






., ,-> <-><!<: 


.o«0 




"/pfo" 


r . - 






... 




■ 


•3 

(S3©' 






^Iffffi 



main bone 
layer 



FIGURE 4. Profiles through the bone bed. Distance from Reference Point 1 to Reference Point 2, approximately 14 
metres. 



The dominant pattern is coarsening upwards, 
with coarse grits overlain by episodic fine silts 
and clays. Several depositional episodes are thus 
represented in the fossiliferous unit. The 
distribution of large basalt cobble is incongruous 
with the surrounding fine-grained sedimentary 
regimes. Apart from obvious association with 
sediments such as in basal erosional polymict 
conglomerate, the basalt cobbles elsewhere are 
randomly distributed. The composition of the 
basalt indicates it is from the nearby outcropping 
flows, and is also comparable in terms of 
deterioration. Smaller basaltic fragments in the 
fossiliferous unit (<2 mm) are severely altered by 
kaolinitization, recognisable only by the black 
specks of ilmenite and magnetite in a clay matrix. 
The weathered basalt clasts also showed vesicles. 

The lowermost sediments of the fossil beds 
contain appreciable basalt, large pieces of 
ferruginized sandstone and intraclasts of dense, 
pure kaolinite. This clearly indicates erosion of 
exposed Werribee Formation (both kaolinitic clays 
and ferruginized sandstones) with input from the 
nearby eroding basalt plain. 

Sediments of the bone bearing unit are mostly 
sand-sized once secondary clay content is 
extracted. The quartz content is of two types: 
orange quartz derived from the ferruginized grits. 
often cemented by iron oxides, and secondly, 
white quartz derived from the purer Werribee 



Formation clays. The latter is hydrothermal quartz 
eroded from Ordovician sediments of the Brisbane 
and Lerderderg Ranges. Rare grains of plutonic 
quartz are also present. The quartz is mostly 
subangular. The only particles exceeding 20 cm 
diameter are basalt cobbles or Diprotodon bones. 
A typical section through the bone bed (Section E, 
Fig. 4) shows that the fossiliferous sediments are 
poorly sorted, platykurtic, near symmetrical 
medium to gravelly sands, with a 41-73% kaolin 
content. 

Hypothetically the ferruginized grits capping 
Hine's Hill eroded back away from the quarry by 
gullying, leaving steep, rilled, erosional faces. The 
base of this scarp has a build up of talus sediment. 
Sediments from the eroding gully talus and 
ephemeral valley stream near the quarry were 
analysed for grainsize and composition (Figs. 5,6). 
The gully sediments are poorly sorted, mesokurtic, 
near symmetrical sandy gravels (Mz= -0.98) on 
top with poorly sorted leptokurtic coarsely skewed 
medium to fine sands below (MZ=1.32-1.92). 
The ephemeral stream sediments are poorly sorted, 
near symmetrical, mesokurtic very coarse sands 
with appreciable (28%) gravel. A sample of 
surficial sediment from the quarry floor was also 
analysed, indicating a poorly sorted, near 
symmetrical platykurtic medium sand. 

In all analyses the clay and silt fractions are 
very low. Sediments closest to the surface are 



100 



J. LONG & B. MACKNESS 



25- 




1 












WT % - 








2 


- 


3 


15- 












■ 




5- 










y^^ 


■ 


/^-\ 


/ 


0.5 


1.5 


2.5 o 


■ i i i i i 




















- 












_ 




- 




4 






5 




6 


- 








- 


/ 




_y\ 


- 








- 


j^y^ 




/~^~^ 




I 


7 




- 


i i i i i i 

/^\ 8 




I 1 1 1 1 ! 










- 


X^ 




y u 




i i 




■ i i i i i 


i i i i i i 



FIGURE 5. Sediment analyses from the bone bed (1-5), basement Tertiary Werribee Formation (6), and recent 
sediments near the fossil site (7-9). All to same scale, equal to axes of graph 1. Dotted lines indicate exclusion of 
secondary fine clays. All MG specimens: 1, -59442; 2, -59443; 3, -59444; 4, -59445; 5, -59441; 6, -59440; 7, - 
59438 (ephemeral tributary stream to the Parwan Creek); 8, -59434 (gully talus sediment); 9, -59439 (surface scree 
from quarry floor). 



coarser and less sorted. Subsurface transportation 
of the kaolinitic fraction apparently occurs rapidly 
after erosion. The steep gradient of the bone bed, 
and the sedimentological analyses, indicate that 
the bones were deposited in an eroding gully and 
covered by surficial scree sediments and 
ephemeral run-off sediments, identical with those 
analysed from the vicinity of the quarry. 

The shallow depth of the fossiliferous unit, 
together with its orientation parallel with the 
topographic surface, suggests it is a relatively 
recent accumulation, accruing over time by 
numerous episodic depositional events. Study of 
the bone surface textures (next section) supports 
the hypothesis that the surrounding sediment 
deposition was not a result of high energy, 
continual processes, but slowly accumulating 
processes. This would allow time for the bones to 
be subaerially exposed for a prolonged period of 
time and weather prior to burial. 



Taphonomy 

Bone orientation data and interpretation 

Stereographic projections, equal area net, 
(Phillips 1971) of bone long axis (poles to bones) 
for the 1978-1979 excavations revealed no overall 
trends (Fig. 1). When proximal (upslope end), 
central and distal sections of the bone bed were 
examined separately a weakly defined trend was 
seen for the distal end of the main Diprotodon 
concentration as bimodally directional. This 
indicates that the highest degree of hydraulic 
sorting occurred at this end, that is the furthest 
downslope area from the proximal concentration 
of skeletons. The information is derived from 
small fragments of bones, mostly rib pieces of 
similar size and shape. This suggests that 
although a small degree of current sorting 
occurred here the orientation of bones forming the 



BACCHUS MARSH DIPROTODON 



101 



WT% 








25- 








20- 




A 




15- 




/ 




10- 


1 •-'""" / 


A /^~~i'.. , 
■h\ /■ 

v--v 


\ 

\ 

\ 

\ 

\ 


5- 


/ 
/ 

/ 




\ 
^ ^' — ■ 




2- J 

1 


— r 1 1 1— 


1 r 



■0.5 0.5 1.5 2.5 35 



3" 



FIGURE 6. Comparison of recent sediments (2,3) with 
sediments of the fossil bed (1,4). All MG specimens: 1, 
-59445; 2, -59439 (surface scree, quarry floor); 3, - 
59443; 4, -59438 (ephemeral stream). 

major portion of the bone bed was not influenced 
by hydraulic processes. Fig. 8 shows the overall 
plot of bone positions both horizontally and 
vertically in the bone bed, based on data from the 
1979 dig. 

Voorhies (1969) demonstrated that the bones 
from animals larger than sheep will act 
responsively to given hydraulic regimes, resulting 
in a sorting of bone groups from proximal to distal 
ends of stream flow. 'Voorhies groups' are 
associations of skeletal elements with similar 
hydrodynamic transportational properties. Group 
1 contains vertebrae, ribs, sacra and sterna, and 
are considered the most easily transported, either 
by floating or saltation. Group 2 contains mostly 
limb elements, and group 3 contains the least 
mobile bones such as skulls and mandibles. 
Elements of group 2 move by traction whereas 
group 3 elements are chiefly lag components, 



which remain closest to the site of death. 
Examinations of the distribution of Voorhies 
groups for the Hine's Quarry Diprotodon bed 
shows a concentration of 95% of group 3 bones at 
the proximal end of the bone bed. Most of the 
skull and jaw elements were closely associated. 
Fig. 9 shows a plan of bones from the central part 
of the bonebed reflecting the orientation of bones 
with respect to channel morphology. The 
individuals were very close to one another at the 
final place of deposition (Figs 10,11). 

The roughly sequential array of Voorhies groups 
seen in the bone distribution map suggests only a 
minor degree of hydraulic transportation. The bone 
orientation data and sedimentological evidence 
indicates that ephemeral stream flows, of high 
energy but short duration, carried the partially 
articulated skeletons from the place of death to the 
main burial site. Subsequently erosional screes 
covered the exposed skeletons. The absence of 
large numbers of group one elements (for at least 
22 individual Diprotodon) further supports the 
influence of rapid water flows for short durations, 
carrying most of these lighter bones away from the 
area which was excavated. 

Bone orientations (Fig. 7) are generally 
horizontal to shallow plunging throughout the 
fossil bed, except for one localised concentration 
of steeply plunging associated macropodid bones 
in the north-eastern boundary of the bone bed. 
These bones rested in unstable positions with 
respect to gravity. 




FIGURE 7. Stereoplot (equal area projection) of bone 
orientation from the 1978-1979 excavations. Note 
cluster of shallow dips and lack of bimodalism. 



102 



J. LONG & B. MACKNESS 



The high degree of bone articulation combined 
with their orientation suggests a separate episode 
from that of the original Diprotodon bone 
accumulation. Hypothetical ly a carcass dumped 
into an irregular erosional gully, somewhat like a 
pothole would produce a similar effect. Many of 
the isolated macropodid bones or partial skeletons 
occur at a different depth to the main Diprotodon 
bone horizon (Fig. 8) and are clearly separate 
burial events. 

A contour map of the surface on which the 
bones were deposited was constructed (Fig. 9) 
from the elevations of the lowest bones in the 
main concentration of the bonebed. The channel 
(or channels) in which the bones were buried had 
an irregular morphology, often with V-shaped 
embayments upslope. The channel bottom was 
notched with local low and high spots. When bone 
orientations are studied in view of channel 
morphology it would seem that this factor had the 
strongest influence on the resting positions of 
bones. Bones caught in localised lows or runs 
have stronger preferred orientations according to 
the downstream flow of the channel than those 
dumped upon broad flat areas of the channel. High 
points are invariably devoid of bones. 

Bone orientations are most easily explained in 
terms of channel morphometry, without indication 
of significant current sorting, save for the extreme 



distal area of the bone bed. Sorting into Voorhies 
groups indicates spasmodic, short durations of 
concentrated high flow regimes, possibly flash 
floods washing carcasses down the gully 
channels. The main proximal concentration of 
articulated skeletons did not move far from the 
site of death, as indicated by their fine 
preservation and high degree of articulation. 

The distal end of the bone bed terminates at the 
quarry scarp, and hence a large number of 
Voorhies group 1 elements are presumably 
missing from the excavated area having been 
washed away. The bone distribution map shows 
that there is a high density of bones proximal to 
the truncation of the bone bed, and it can be safely 
assumed that most of the fossil remains of 
Voohries groups 2 and 3 were recovered by the 
excavations. 

Bone preservation and articulation 

Most of the bones have poor preservation, being 
soft and crumbly due to preburial weathering and 
diagenesis. Exposure of the bone before burial 
results in steady decomposition according to 
localised conditions (Behrensmeyer 1978; 
Behrensmeyer et al. 1979). The degree of bone 
surface decomposition ranges from Type 1 
(Behrensmeyer 1978) (seen only in the proximal 





M 1 












^^ 


""■"*: 




REF. PT. 2 


[."] 






' : ';"- ^^^ 


-~- """ "^ — -- 


+ 


y' -\ macropod remains 






— — 










""" REF. PT. 1 






\ ■ 


\ 


-1 




main bone layer 










-2 




\ / 








"X 

'•-.■■■■■\ 


-3 




\ / 












random occurrences 






r-i 


-M 













FIGURE 8. Distribution of bones from the 1978-1979 excavations. Top, plan view showing proximal channel 
outline. Bottom, depth of bone distribution. 



BACCHUS MARSH DIPROTODON 



103 



region of the bone bed) to almost completely 
decayed pieces of bone, where the surface has 
been lost, showing only the spongy trabecular 
bone. Overall, most bones show surface flaking 
and cracking parallel to bone fibre, with a small 
degree of exfoliation of the outer surface, 



corresponding to Types 2 to 3 of Behrensmeyer's 
scale (Fig. 13: 3,4). Comparison with the results 
from the Amboseli basin of Africa suggests that 
such bone has undergone prolonged exposure to 
arid climatic conditions. 
Moisture accelerates bone decomposition, and 



130 




FIGURE 9. Channel morphometry shown in relation to bone orientation. Contours plotted by lowest bone depth from 
main concentration, bone orientations and preservation correctly shown. 



104 



J. LONG & B. MACKNESS 








... !-..„...- 








■,'::*!p| 






: 



FIGURE 10. Photograph from the 1973 excavation at the extreme proximal portion of main bone bed. Note excellent 
preservation, dense concentration and partial articulation of bones. Photo courtesy Ian Stewart. 




FIGURE 1 1 , Partially articulated skeleton in the proximal area of the bone bed, 1973 excavation. Photo courtesy Ian 
Stewart. 



BACCHUS MARSH DIPROTODON 



105 




FIGURE 12. Bone preservation from the main bone bed. Seventeen Diprotodon femora showing degree of 
completeness and approximate state of surface condition. 



decay of skeletons in temperate to tropical 
environments results in different styles of bone 
deterioration (Behrensmeyer 1975). Separate teeth 
of Diprotodon from the distal area of the bone bed 
show fractured roots and streaking of the enamel 
(Fig. 13: 1,2). Root fracturing is due to trans- 
portation, with significant preburial exposure 
causing enamel streaking and eventual splitting 
(Behrensmeyer 1978). 

In general, the bones are recovered in variable 
states of preservation and degrees of articulation. 
The most proximal end of the bone concentration 
(1973 dig) shows a high degree of articulation, 
and relatively good preservation of bone surface 
(Figs 10,11). Articulation is not complete in any 
instance, the most complete skeletal remains 
comprise of skull, mandibles, vertebral column, 
parts of the limbs, shoulder girdle, pelvis and feet, 
not all articulated, but in close proximity. Skulls 
in the proximal area of the bone bed were often 
turned over, and one mandible was upside down 
resting in a steep position. Limb bones were 
always lying flat, as were scapulae and pelves. 
The strongly curved nature of the articulated 
vertebral column (Fig. 11) could indicate bending 
due to ligament contraction following drying of 
the carcass. The high number of skulls and 
mandibles recovered from this area, and the high 
degree of postcranial articulation, relative to that 
seen in the distal end of the bone bed suggests 
close proximity to the place of death. 

The distal end of the bone bed is composed of 



non-articulated limb and rib pieces, in variable 
state of preservation. Skulls and jaws were 
concentrated at the proximal end of the bone bed. 
One isolated skull was found very close to the 
distal end of the bone bed, and had presumably 
rolled down. Sections of articulated Diprotodon 
vertebral column were found only in the proximal 
half of the 1978-1979 excavation. 

To conclude, the overall degree of articulation 
shown throughout the bone bed is not indicative 
of a natural trap or catastrophe in which animals 
are rapidly entombed whole (e.g. bogging, pitfall) 
but suggestive of mass mortality followed by 
exposure and some degree of transportation. At 
the proximal end of the bone bed some part of the 
skeletons were buried rapidly and show good 
surface preservation. The majority of the carcasses 
were disarticulated and moved slightly downslope. 

Condition of the bones 

Since the sediments of the bone bed were rich 
in kaolinitic clays, the diagenetic environment was 
basic (a solution of distilled H 2 with sediment 
MG 59443 was slightly basic, pH=7.5, although 
when made into a mud slurry with little water 
pH=9). High porosity of the grits in the bone bed 
enable concentrated basic condition for bone 
decomposition. 

Microprobe analyses of fossil bone and tooth 
fragments from the bone bed show a slight 
increase in calcium oxide content, a decrease in 



106 



J. LONG & B. MACKNESS 




FIGURE 13. Bone preservation. 1-2, Isolated Diprotodon molars (no numbers) showing enamel streaking and 
splitting, with transportational abrasion to roots; 3, Highly weathered splinter of macropodid bone, showing parallel 
surface exfoliation, splitting and fracturing; 4, Macropodid jaw from the distal end of the main Diprotodon 
concentration. Note excellent preservation of posterior molars with splintering of premolar and anterior two molars. 
The ramus shows preburial exposure cracking and flaking, with transportational breakage to diastema and ascending 
ramus. 



phosphoric oxide P 2 5 with chloride salts added at 
2-3% (relative to probed samples of modern 
Macropus bone and teeth). This indicates that the 
bones lacked mineralisation because of constant 
leaching or disappearance of available calicum 
oxide. Drying and wetting of the clays around the 
bones may have caused diagenetic fracture by 
swelling and compaction as host rock volume 
changes. Many bones had a coating of a black 
mineral presumably manganese salts. 

Very few bones were preserved in their entirety, 
although those that were show only minor 
abrasive features such as rounding of edges. 
Vertebrae with entire neural arches typify this 
state, although they show gentle rounding of 
extremities. In a study of carnivore gnawing 
patterns on recent and Pleistocene mammals, 
Haynes (1980) documented the individual patterns 
of damage incurred upon each skeletal element 
when wolves utilised carcasses of medium to 
large-sized herbivores. Some of the Bacchus 
Marsh Diprotodon bones demonstrate similar 
damage patterns to those of bison scavenged upon 
by wolves. 



Out of 28 femora examined only two were 
completely preserved without wear or breakage, 
apart from very slight abrasion of the greater 
trochanter (P 150114, and #94; Fig. 12). Three 
specimens showed good preservation of the 
trochlear area but had damaged greater trochanters 
or were lacking the femoral head. Light utilisation 
of bison femora by wolves results in damage to 
the greater trochanter only, followed by damage to 
the distal condyles and removal of the femoral 
head during heavy utilisation. The shaft may not 
be damaged as the end of the bone is softer, 
enabling easier access to marrow. 

The majority of Diprotodon femora recovered 
from the site are missing both femoral head and 
trochlear region (Fig. 12). Fractures on the distal 
end of their shafts are screw type (Haynes 1980), 
suggesting transportation breakage. If gnawing 
had occurred prior to transportation then rounding 
of the edges, through abrasion, could have 
smoothed the jagged edges produced through 
carnivore action. Evidence of preburial scavenging 
is seen on some of the tibiae which hypothetically 
show gnawed off proximal heads (e.g. #67, #164, 



BACCHUS MARSH DIPROTODON 



107 



#150, P150203). 

Haynes (1980) shows articulated humeri with 
light carnivore gnawing, such as damage to the 
distal end, rather than the commonly gouged out 
and broken proximal epiphyses. The Diprotodon 
humeri showed much damage to the proximal 
epiphyses (#47, #157, #162, #257, #424, 
P151889, #218). Although no similar studies have 
been made on Australian carnivores, 
contemporaneous animals such as Thylacoleo 
were of suitable size and have left fossil evidence 
of predation on Diprotodon and other animals 
(Horton & Wright 1981; Runnegar 1983). 
Reptilian scavengers such as the giant varanid 
Megalania should also not be ruled out as 
contributing to post mortem bone damage. 

The main bone concentration contains all the 
Diprotodon bones found, with only few additional 
macropodid bones, rodent remains and a lizard 
jaw. All other bones were found at various depths 
throughout the sediment, not confined to any 
single horizon or lithology (Fig. 8). A rodent 
maxilla was found in association with a partial 
macropodid skeleton in the distal part of the site. 

In summary, both the state of bone preservation, 
bone fracture patterns and the distribution of taxa 
throughout the sediment suggest that some bones 
were transported and subaerially exposed longer 
than others. Different burial events are responsible 
for the main Diprotodon element concentration 
than for individual macropodid burials. 

Bone settling velocities 

Behrensmeyer (1975) formulated a method for 
calculating bone settling velocities in which bones 
were equilibrated to quartz pebbles sizes. For 
bones the size of Diprotodon, large quartz 
spheroids would be necessary. For example, a 
hippopotamus premolar has a quartz spheroid of 
13.8 mm diameter as its hydraulic equivalent. 
Ribs of cows (Diprotodon size) require only 3.1 
mm quartz equivalents. The flow of water moving 
the coarser quartz sands and grits of the bone bed 
may have just reached high enough energies to 
move ribs, but did not reach velocities capable of 
transporting larger skeletal elements by 
themselves. For vertebrae of about 29 cc volume 
(macropodid size) the hydraulic equivalent is a 
quartz spheroid of 4.4 mm diameter. 

Most bones from the site have a much higher 
hydraulic stability than their entombing sediments, 
indicating close proximity and little transportation 
from the site of death. Movement to the distal area 
of the bone bed could have occurred either by 



flotation of carcasses prior to decay, or have been 
assisted by the high slope gradient of the exposed 
channel, thus requiring lower flow regimes to 
achieve higher degrees of transportation. 



Biological Data - population structure 

At least 22 individual diprotodontids (based on 
numbers of right femora), all probably 
representing a single species of Diprotodon, were 
recovered from the site. A number of macropodids 
were also present as well as a variety of smaller 
mammalian taxa including rodents, dasyurids and 
peramelids. Fossils of reptiles have also been 
recovered. 

It is clear from the tooth eruption and wear 
patterns of the Hine's Quarry Diprotodon that 
most were adults. Of the sixteen mandibles (Fig. 
14) examined, most had all molars erupted and 
the majority of specimens lacked wear on both 
lophs of the fifth molar. Aged and juvenile 
individuals are absent. 

The taphocoenosis (death assemblage) of the 
deposit is biased towards similarly aged 
individuals and excludes the very young and the 
very old. Such a selection could represent random 
sampling (normal rarity of juveniles or aged) or a 
biased selection of the strongest or most active 
animals. Similar age structure is described by 
Newsome (1965) in studying the reproduction of 
female red kangaroos in drought conditions. He 
noted that fecundity decreases during drought 
conditions and that adults in their prime constitute 
the dominant surviving group. The macropodid 
jaws from the deposit include at least two 
juveniles (e.g. MV-150120), although it must be 
pointed out that these apparently represent 
separate burial episodes from the main 
Diprotodon concentration. 



Discussion 

Both the sediments of the bone bed and the 
distribution and orientation of the bones indicate 
that deposition occurred at the proximal end of an 
ephemeral stream system. The system envisaged 
would be at the foot of a series of gullied rilles 
running from the top of the hill (straighter at the 
proximal end) to the creek (increasing sinuosity 
distally). Evidence for this is seen by the 
discontinuous nature of bedding with lenticular 
and festoon cross-stratification. The latter 
indicates fluctuating sedimentary regimes and 



108 



J. LONG & B. MACKNESS 



moderate to lower flow regime (Dane-Picard & 
High 1973). 

Thin wisps of almost pure clays and silts, 
occurring randomly throughout the bone bed are 
interpreted as depressions in the exposed surface 
where clays settled out of puddles following 
intense rains. This phenomenon was observed to 
occur in the quarry during fieldwork when 1-2 cm 
kaolinitic layers formed at the bottoms of puddles 
from overnight rain. It is because of the 
impermeable nature of the kaolinitic basement that 
the erosion of clays occurs. The presence of large 
basalt cobbles throughout the bone bed can be 



1 






U 00 coo 

GC coco 
( (U0o<*> 



COGOocd 




explained by collapse and rolling of the nearby 
eroding basalt plain, hence the irregularity of the 
cobble size and distribution. 

A close relationship is seen between sediments 
containing the bones and the modern sediments of 
the area. Mass percentage versus grain-size for 
some of these are identical. Sediment samples MG 
59445 and MG 59439 (Fig. 5, graphs 4 and 9) are 
both poly modal with 4 peaks each, all 
corresponding to 1 phi size, most within 0.5 phi. 
Highest peaks are reached as 2.5 phi for 59439 
and 3 phi for 59445. This demonstrates that the 
sediment containing the bones is little different 



&8 




J coco 




a (iao 




11'! 



^HUd(fO 



DOG 






ffOQGOo 




(Ttfoo 



16(KJ (&<£<?* 



FIGURE 14. Sketches of tooth eruption patterns for sixteen individual Diprotodon mandibles. Where both sides were 
present the best preserved tooth is shown. 1, P151801; 2, P151802; 3, P150293; 4, P32223; 5, unnumbered; 6, 
P15106; 7-8, unnumbered; 9, P150017; 10, P151804; 11, #492; 12, P150062; 13, P150298; 14, P150299; 15, #25 
(1973); 16, unnumbered. 



BACCHUS MARSH DIPROTODON 



09 



from the surface wash scree on the quarry floor 
(MG 59439, Fig. 6, graph 2) which was derived 
from erosion of both ferruginized grits and purer 
sandy clays of the Werribee Formation. 
Comparison with ephemeral stream sediments 
near the quarry (MG 59438, Fig. 6, graph 4) 
indicates episodes of higher flow regime. Both 
MG 59438 (modern ephemeral stream) and MG 
59443 (bone bed; Fig. 6, graph 3) are unimodal 
with closely spaced highest peaks at -0.5 phi (MG 
59438) and O phi (MG 59443). Maxima for these 
samples are 16.7% and 13.2% respectively. This 
suggests that the sediment of the bone bed may 
have had a slightly lower energy of deposition. 

The sediments entombing the bones signify a 
stream or gully system of generally low flow 
regime and fluctuating deposition. Erosional 
screes are intercalated with spasmodic higher flow 
regime channel sediments. Irregular channel 
morphometry and random bone orientation further 
supported by the steep channel gradient suggest a 
juvenile hydraulic system buried the carcasses. In 
summary the evidence does not favour a regular 
fluviatile channel environment as first suspected, 
but an ephemeral erosional system. 

The sedimentological data favours an arid 
climate at the time of deposition. Minimal 
predator influence corroborates the idea that aridity 
may have caused the death of a Diprotodon herd. 
Considering that the fossil-bearing sediments are 
close to the modern ground surface, the 
topography at the time of the event was probably 
not unlike the present condition. The eroded scarp 
of the ferruginized grits on top of Hine's Hill and 
the basalt would have been closer to the fossil site. 
The bone bed morphology is comparable to an 
eroding gully rille presently exposed above the 
fossil site, and the fact that the bones are 
channelled along a similar direction to the existing 
hill topography supports the recency of the event. 

Ephemeral hydraulic regimes also support the 
aridity hypothesis, which would suggest an age 
for the site at the most recent late Pleistocene 
glacials (Bowler 1982). The phylogeny of the 
Diprotodontidae suggested by Stirton et at. (1967) 
restricted the genus Diprotodon to the Pleistocene, 
although the genus is also known from the late 
Pliocene Kanunka Local fauna (Tedford, Williams 
& Wells, 1986). 



The scenario suggested by the combined data is 
that the herd of young adult Diprotodon expired 
from a period of severe aridity and drought. The 
carcasses were subsequently moved downslope 
after scavenging and extensive subaerial 
decomposition. The highly basic nature of the 
entombing sediment pore fluids caused a rapid 
deterioration of the fossil bones. The evaporation 
of ground waters during intense aridity resulted in 
the secondary infilling of kaolinitic clays in the 
pore spaces of the bone bearing sediments. This 
infilling was assisted by erosion of clay rich 
sediments from the surface which seeped 
downwards between pore spaces in following 
seasons of rainfall. 



Acknowledgments 



The initial field work for this study was undertaken as 
a third year geology project (J.L.) in the Department of 
Earth Sciences at Monash University in 1979. It was 
suggested by Tom and Patricia Vickers-Rich who 
provided useful guidance. Tom Rich also provided the 
first set of bone orientation readings from the 1978 field 
season. Tim Flannery, Rob Glenie, Ian Stewart, Thomas 
Darragh, Dominic Williams (now deceased), Arthur 
Day, John Clemens, Greg McNamara and Robert Baird 
provided assistance and discussion on many aspects of 
the work. Ian Stewart made a series of panoramic 
photographs from the 1973 field season available for 
study. John Hamilton (CSIRO Mineral Physics, Port 
Melbourne) facilitated the use of a scanning electron 
microscope and DTA equipment for study of clay 
samples and bone surface condition. Mr. A Taylor kindly 
provided accommodation during field work, and the 
Hine family permitted excavation of the site by halting 
quarry work. 

The study of the Bacchus Marsh Diprotodon material 
was supported in part (B.M.) by an ARC Program Grant 
to Michael Archer; a grant from the Department of Arts, 
Sport, the Environment, Tourism and Territories to 
Michael Archer, Sue Hand and Henk Godthelp; a grant 
from the National Estate Program Grants Scheme to 
Michael Archer and Alan Bartholomai; and grants in aid 
to the Riversleigh Research Project from Wang Australia, 
ICI Australia and the Australian Geographic Society. 

Excavation of the Bacchus Marsh Diprotodon material 
was financed by the Council of the Museum of Victoria. 
The preparation of this material was paid for by grants 
to Thomas H. Rich from ARC. 



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the Museum of Comparative Zoology, Harvard 146: 
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no 



J. LONG & B. MACKNESS 



ecologic information from bone weathering. 
Paleobiology 4: 150-162. 

BEHRENSMEYER, A. K., WESTERN, D. & BOAZ, D. 

E. D. 1979. New perspectives in vertebrate 
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BOWLER, J. M. 1982. Aridity in the late Tertiary and 
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COOKSON, I. C. 1957. On some Australian Tertiary 
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DANE-PICARD. M. & HIGH, L. R. 1973. 'Sedimentary 

structures of ephemeral streams: development in 
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FENNER, C. 1918. The physiography of the Werribee 
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FOLK, R. L. 1961. 'The Petrology of Sedimentary 
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GILL, E.D. 1964. Rocks contiguous with the basaltic 
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Royal Society of Victoria 77 : 3 3 1 -3 55 . 

HAYNES, G. 1980. Evidence of carnivore gnawing on 
Pleistocene and Recent mammalian bones. 
Paleobiology 6: 341-5 1 . 

HORTON, D. R. 1976. Lancefield: the problem of proof 
in bone analysis. The Artefact 1: 129-143. 

HORTON, D. R. & SAMUEL, J. 1978. Paleopathology 
of a fossil macropod population. Australian Journal 
of Zoology 26: 279-92. 

HORTON, D. R. & WRIGHT, R. V. S. 1981. Cuts on 
Lancefield bones: carnivorous Thylacoleo, not 
humans, the cause. Archaeology in Oceania 16: 73- 
80. 

NEWSOME, A. E. 1965. Reproduction in natural 
populations of the red kangaroo, Megaleia rufa 
(Demarest), in Central Australia. Australian Journal 
of Zoology 13: 735-759 

PHILLIPS, F. C. 1971. 'The use of stereographic 
projection in structural geology'. Edward Arnold: 
London. 90 pp. 

PLEDGE, N. S. 1993. A century of Callabonna research. 
Conference on Australasian Vertebrate Evolution, 
Palaeontology and Systematies. April 19-21 Adelaide. 
(Abstract) 

RAHMAN, A. & MCDOUGALL, I. 1972. Potassium- 
argon dates on the Newer volcanics of Victoria. 



Proceedings of the Royal Society of Victoria 85: 
61-9. 

RICH, T. H. V. 1976. Recent fossil discoveries in 
Victoria; five late Cenozoic fossil marsupial sites in 
Victoria; a progress report. Victorian Naturalist 93: 
198-206. 

RUNNEGAR, B. 1983. A Diprotodon ulna chewed by 
the marsupial lion, Thylacoleo carnifex. Alcheringa 
7: 23-25. 

SUMMERS, H. S. 1923. The Geology of the Bacchus 
Marsh and Coimadai District. Proceedings of the 
Pan-Pacific Science Congress (Australia) 2: 1632- 
48. 

TEDFORD, R. H. 1973. The Diprotodons of Callabonna. 
Australian Natural History 17: 349-54. 

SCOTT, H. H. 1915. A monograph on Nototherium 
tasmanicum (Genus - Owen: Sp. nov.). Tasmanian 
Department of Mines. Geological Survey Record 4: 
1-47. 

STIRTON, R. A., WOODBURNE, M. O. & PLANE, M. 
D. 1967. A phylogeny of the Tertiary Diprotodontidae 
and its significance in correlation. Bulletin of the 
Bureau of Mineral Resouces, Geology and 
Geophysics, Australia 85: 149-160. 

TEDFORD, R. H. 1993. Lake Callabonna: "Veritable 
necropolis of gigantic extinct marsupials and birds" 
Conference on Australasian Vertebrate Evolution, 
Palaeontology and Systematies. April 19-21 Adelaide. 
(Abstract). 

TEDFORD, R. H., WILLIAMS. D. G. & WELLS, R. T. 
1986. Late Eyre and Birdsville Basins: Late Cainozoic 
sediments and fossil vertebrates. In R. T. Wells & R. 
A. Callen (eds.), 'The Lake Eyre Basin - Cainozoic 
Sediments, Fossil Vertebrates and Plants, Landforms, 
Silcretes and Climatic Implications' . Australian 
Sedimentological Group Field Guide Series No. 4, 
Geological Society of Australia, Sydney. 

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Geology of the Brown Coals of Victoria. (Part 3.). 
Mining and Geology Journal 4(2): 41—63. 

VAN HUET, S. 1993. The Lancefield megafaunal site, 
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Evolution, Palaeontology and Systematies. April 19- 
21 Adelaide. (Abstract) 

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dynamics of an early Pliocene vertebrate fauna, Knox 
County, Nebraska. University of Wyoming 
Contributions to Geology, Special Papers. 1: 1-69. 

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Cainozoic volcanic rocks of eastern Victoria, 
Australia. Journal of the Geological Society of 
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Society of Victoria 11: 273-327. 



A NEW FOSSIL WALLABY (MARSUPIALIA ; MACROPODIDAE) FROM 

THE SOUTH EAST OF SOUTH AUSTRALIA 

7. A. McNamara 



Summary 

A new wallaby, Congruus congruus gen. et sp. nov., is described, from a cave fill presumed to be of 
late Pleistocene age. While agreeing in some characters with many other macropodine genera, it 
most resembles Prionotemnus and Protemnodon. 



A NEW FOSSIL WALLABY (MARSUPIALIA; MACROPODIDAE) 
FROM THE SOUTH EAST OF SOUTH AUSTRALIA. 



J. A. McNAMARA 



McNAMARA, J. A. 1994. A new fossil wallaby (Marsupialia; Macropodidae) from the South 
East of South Australia. Rec. S. Aust. Mus.: 27(2) 1 1 1-1 15. 

A new wallaby, Congruus congruus gen. et sp. nov., is described, from a cave fill presumed 
to be of late Pleistocene age. While agreeing in some characters with many other macropodine 
genera, it most resembles Prionotemnus and Protemnodon. 

J. A. McNamara, South Australian Museum, North Terrace, Adelaide, South Australia 5000. 
Manuscript received 17 June 1993. 



Mammal faunas from the cave fills of the 
Naracoorte area have been reported by Williams 
(1980), Wells et al. (1984) and Pledge (1990). 
These caves have yielded big samples of medium 
to large macropods, but, so far, few new forms 
have been reported, notably Sthenurus maddocki 
(Wells and Murray 1979) and Sthenurus 'P17250' 
(Prideaux et al., in press). Just as the highly 
distinctive but apparently rare vombatid Warendja 
is known from four specimens from only two 
localities, so it might be expected that, as 
collections grow and are examined critically, 
specimens of rarer forms will be discovered. 

The unique type specimen was compared 
directly with all specimens available in the fossil 
and modern mammal collections of the South 
Australian Museum. It was prepared by hardening 
with dilute Bedacryl after it was decided that it 
was too delicate to be safely handled; nor could a 
patina of fine sand, cemented with lime, be 
removed without losing bone. The teeth of the left- 
hand side were hand cleaned. 

Tooth numbers follow Archer (1978). The 
composition of the Macropodinae used is that of 
Flannery (1989) but without Hadronomas 
(Murray, 1991). 

Fig.2 contains an orientation symbol consisting 
of an arrow pointing to the anterior and an 
upturned U representing the tongue or lingual 
side. 



Systematics 

Family MACROPODIDAE Gray 1821 
subfamily MACROPODINAE Thomas, 1888 

Congruus congruus gen. et sp. nov. 



Holotype 

P33475 (registered in the palaeontological 
collection of the South Australian Museum), a 
nearly complete adult skull with ?\ M 2 , to M 5 
both I 2 and left P, missing anterior part of left 
nasal, the right zygoma, part of right temporal and 
mastoid. The incisors show moderate wear. P 2 is 
unworn. The molars grade from the moderately 
worn M 2 to the unerupted M 5 . 

Locality 

S.O.S. cave (5U132) just south of Naracoorte in 
the South East of South Australia. 

Age 

Late Pleistocene by faunal association. 

Etymology 

From the Latin for agreeable or harmonious. 
Gender masculine. 

Diagnosis 

Congruus agrees in many of its character states 
with other members of the Macropodinae, but 
more closely resembles Protemnodon, 
Prionotemnus, Kurrabi, Wallabia and Macropus. 
It can be distinguished from the other 
macropodine genera by many characters, including 
lack of canine, long diastema, higher-crowned 
molars, entire palate and size. 

Congruus is distinguished from Protemnodon 
by possessing a deflected rostrum; a P 2 shorter 
than most molars (M 3 , M 4 , M 5 ); a rather small 
masseteric process, not extending down to the line 
of the alveolar margin; and by lacking a large 
labial groove on P. 

It differs from Prionotemnus in possessing a 
more anterior placement of the infraorbital 
foramen; a less distinctly grooved P 2 , with a wider 



112 



J.A.McNAMARA 




FIGURE I. Congruus congruus holotype skull P33475, x 0.75. 



NEW FOSSIL WALLABY 



113 



TABLE 1. Measurements (mm) of skull of Congruus 
congruus (P33475, holotype). 



Description 



mm 



maximum length sans teeth 160.0 

maximum height 59.4 

maximum width of frontals 50.3 

maximum width of nasals 36.7 

length of diastema between alveoli 39.9 

palate width at mid-diastema 20.8 

rostrum width at mid-diastema 22.6 

maximum width of occiput 21.8 

width of postorbital constriction 3 1 .2 

height of premaxilla 30.7 

longitudinal basin; a V 1 shorter than M 5 ; higher- 
crowned molars with longer, more procumbent 
anterior cingula; a forelink on M 2 and M- 3 ; and 
well-developed posthypocristae. It differs in 
lacking the deep groove well within the labial 
surface of P. 

From Kurrabi it may be distinguished by its 
relatively short P 2 ; shorter, more anterior 
masseteric process; less elongate molars, having 
an oblique posthypocrista and lacking a distinct 
posterior fossette. 

From Wallabia, Congruus is distinguished by 
its lacking a prominent labial groove on P; well 
developed labial crests on all molars; and an 
anterior nasal spine. It is further distinguished by 
possessing procumbent incisors, an entire palate, 
a P- 3 shorter relative to the molars, with a smooth 
longitudinal basin; and by having the anterior 
portion of the brain case less constricted. 

It differs from Macropus in lacking the distinct 
anterior (postorbital) constriction of the brain case; 
a groove on the labial surface of P; a developed 
posterior fossette on the molars; and the inflation 
of the nasal part of the rostrum, relative to the 
anterior palate. Congruus is distinguished from 
Macropus by possession of more procumbent 
incisors; an entire palate; a well-developed ovale 
crest; oblique posthypocristae; and in having the 
loph-crests less bowed or preparacristae less 
developed. 

Description 

The skull, in general aspect, is lightly built, with 
a relatively large brain case, comparing in its 
gracility, with many living Macropus species. 

From the side the skull presents a generally flat 
dorsal profile. The incisors and premaxillae are 
procumbent. No anterior nasal spine is present but 
in this position the premaxillae are smooth and 
depressed. The deep rostrum is arched dorsally 
and is near the plane of the flattened, somewhat 



depressed frontal bones and the slightly raised 
frontoparietal region. The parietals decline 
towards a slight lambdoidal crest. A large 
diastema reveals a palate declining from a rather 
level cheek tooth row. The infraorbital foramen is 
above the anterior half of P 2 . The orbit appears 
relatively small and the zygomatic arch, light and 
shallow. The masseteric process is opposite the 
protoloph of M 4 , and small, not reaching the level 
of the alveolar margin. 

From above, the rostrum tapers evenly forward 
from the lacrimals, and is flat sided without lateral 
inflation of the nasal part. The nasals have a 
broad, fairly straight contact with the frontals, on 
a line with the lacrimal foramina. The frontal 
bones are broad and flat, inflated laterally, above 
the orbit, and slightly depressed, centrally, on their 
common suture. The anterior part of the brain case 
is not greatly constricted postorbitally as it is in 
many macropodine genera. There is no sagittal 
crest and the temporal foramen is small. 

From below, the incisive foramina are small. 
There are no canines. The palate, anterior to the 
cheek teeth, nearly equals the width of the rostrum 
above. The palate appears to have been entire. 
Pterygoid cavities, appearing small, have their 
lateral borders formed by prominent anterior- 
directed ovale crests. The alisphenoid bulla is 
slightly inflated and the auditory process is short. 

Teeth. 

I 1 is unknown but the alveolus is slightly larger 
than that of P, rounded, narrowed ventrally and 
not much compressed laterally. 

P shows no sign of an occlusal groove, perhaps, 
due to wear. The corresponding structure on P is 
attenuated and may indicate that the groove was 
much reduced or lacking on P. A broad shallow 
groove runs parallel and just anterior to the 
posterolabial edge, of P, which is raised and ridge- 
like. 

P has a narrow and shallow occlusal groove 



TABLE 2. Measurements (mm) of upper cheek teeth of 
Congruus congruus (P33475, holotype) 





Length 


Anterior 


Posterior 






width 


width 


p* 


9.8 


4.6 


5.0 


M 2 


8.6 


6.9 


7.1 


M 2 


10.6 


8.1 


7.8 


M 4 


11.5 


8.6 


8.2 


M 4 (estimated) 


10.4 


8.9 


8.0 


P*-M* 


37.7 


— 


- 


V 1 - M 1 (estimated) 


48.1 


- 


- 



114 



J. A. McNAMARA 




FIGURE 2. Congruus congruus holotype P33475, stereopair of left cheek teeth, x 1.5. 



which opens near the posterior edge of the labial 
surface so that the small lingual crest is barely 
visible and the groove so formed is barely visible 
on the labial surface. I have interpreted a pit, 
midway along the posterolabial edge of this tooth, 
as pathological but of localised effect and not 
associated with any general distortion of the crown 
- a view supported by the alveolus of the right P 
which indicates a very similar tooth. It is probable 
that all the incisors were high crowned, and that, 
their relative sizes were I 1 >I 2 >P. Their combined 
outline in occlusal view was probably U-shaped. 

P 3 is a little longer than M i . Its outline is not 
concave labially. There is a prominent labial crest 
with anterior and posterior cusps, with three 
indistinct cuspules and ridgelets between. There is 
a prominent posterolingual cusp, lower than the 
posterior cusp, and connected to it by a ridge, 
behind which is a small shallow posterior fossette. 
From the posterolingual cusp, a lingual cingulum 
runs forward, and is notched at one third of the 
tooth length, then is somewhat raised, before 
ending, almost opposite the anterior cusp. This 
cingulum, together with the main crest, forms a 
smooth longitudinal basin. 

The upper molars are plain and quadrate, 
becoming more elongate and increasing in length 
from M 2 to M 4 and probably M 5 . The anterior 
cingulum is broad and shelflike with a shallow 



basin between it and the protoloph to which it is 
connected by a preparacrista. A low forelink is 
present on M 2 and M 2 but not visible on M- or M 5 . 
The width and length of the anterior cingulum 
increases from M 2 to M 5 . The anterior width of M 2 
is less than its posterior width but near equal in 
the other molars. The lophs have their apical width 
nearly equal to the basal width, not markedly 
narrower as in many Macropus and Kurrabi. A 
midlink is formed just lingual to the middle of the 
transverse valley by the postprotocrista, there is a 
small contribution from the metaloph. A shallow 
basin is formed in the transverse valley by an 
extension of the postparacrista on M 2 and M\ but 
not, M 4 and M 5 . The posterior face of the metaloph 
is rather plain and flattened, with a distinct 
posthypocrista rising obliquely to join the base of 
the metacone, where a small groove separates it 
from a much less distinct, near vertical, 
postmetacrista. Together they do not form a 
centrally placed fossette, seen in many forms, 
including Macropus, Protemonodon* Thylogale, 
Wallabia and Onychogalea. A much reduced 
postlink is discernible on M-. 



Discussion 
The age of this material is inferred from its 



NEW FOSSIL WALLABY 



115 



association with Thylacinus and Sthenurus 
PI 7250 material with very similar preservation. 
This Sthenurus is known from the dated deposits 
of Victoria Cave (Wells et al. 1984) and the 
Henschke Fossil Cave (Pledge 1990), Naracoorte, 
and Green Waterhole (Newton 1988), Tantanoola. 

Discovery of further specimens, particularly 
those with the lower dentition and deciduous 
cheek teeth, should add to the understanding of 
this form and, in particular, clarify its relationship 
to Prionotemnus. 

One's attention is drawn to the prominent naso- 
frontal development of this species, which is 
presumably an autapomorphic character. 
Comparisons can be made with the similar 
structures found in Onychogalea unguifera and, if 
its function were known, it would allow 
conclusions about the functional adaptation of the 
fossil form. 

In Congruus a combination of many primitive 



macropodine features seem to be the foundation, 
overlain by apomorphy in the form of the whole 
skull giving it the general aspect of a modern- 
looking kangaroo. While this is adding to the 
increasingly large puzzle that is macropod 
phylogenetics, it is to be hoped that future study of 
this specimen will shed light on the major 
pathways followed by this family in its great 
evolutionary flowering through the latter half of 
the Neogene to the present. 



Acknowledgments 

I wish to thank Mr A. D. (Tony) Colhoun for 
recovering the material and lodging it with the South 
Australian Museum. Thanks are also due to: N. S. 
Pledge, and G. Prideaux, for reading a draft of this 
paper; D. Van Weenen, for typing; and T. Peters for 
photography. 



References 



ARCHER, M. 1978 The nature of the molar- pre molar 
boundary in marsupials and a reinterpretation of the 
homology of marsupial cheek-teeth. Memoirs of the 
Queensland Museum 18(2): 157-164. 

FLANNERY, T. F. 1989. Phylogeny of the 
Macropodoidea; a study in convergence Pp. 1-46. In 
'Kangaroos, Wallabies and Red Kangaroos.' G. Grigg, 
P. Jarman & I. Hume. Surrey Beatty & Sons Pty Ltd., 
Sydney New South Wales. 

MURRAY, P. 1991. The sthenurine affinity of the Late 
Miocene kangaroo, Hadronomas puckridgi 
Woodburne (Marsupialia, Macropodidae).v4/c/i£/-m£a 
15(4): 255-283. 

NEWTON, C. A. 1988. A Taphonomic and 
Palaeoecological Analysis of the Green Waterhole 
(5L81), a submerged Late Pleistocene Bone Deposit 
in the Lower South East of South Australia. Thesis for 
Honours Degree of Bachelor of Science at the Flinders 
University (School of Biological Sciences). 



PLEDGE, N. S. 1990. The Upper Fossil Fauna of the 
Henschke Fossil Cave, Naracoorte, South Australia. 

Memoirs of the Queensland Museum 28(1): 247-262. 

PRIDEAUX, G. J. & WELLS, R. T. A new extinct 
kangaroo form southeastern Australia. In prep. 

WELLS, R. T. MORIARTY, K. & WILLIAMS, D. L. G. 
1984. The fossil vertebrate deposits of Victoria Fossil 
Cave Naracoorte: an introduction to the geology and 
fauna. The Australian Zoologist 21(4): 305-333. 

WELLS, R., MURRAY, P. 1979. A new sthenurine 
(Marsupialia, Macropodidae) from southeast South 
Australia. Transactions of the Royal Society of South 
Australia 103(8): 213-219. 

WILLIAMS, D. L. G. (1980). Catalogue of Pleistocene 
vertebrate fossils and sites in South Australia. 
Transactions of the Royal Society of South Australia 
104(5): 101-115. 



CETACEAN FOSSILS FROM THE LOWER OLIGOCENE OF SOUTH 

AUSTRALIA 

Nevilles. Pledge 



Summary 

A single, damaged cetacean tooth from the upper part of the Buccleuch Formation of the Murray 
Basin dates from the early Oligocene. It has been compared with Eocene archaeocetes and with late 
Oligocene cetaceans. Similarities are seen with Mammalodon colliveri and Metasqualodon 
harwooddii, but no definite assignment can be made. 



CETACEAN FOSSILS FROM THE LOWER OLIGOCENE OF SOUTH AUSTRALIA 



NEVILLES. PLEDGE 



PLEDGE, N. S. 1994. Cetacean fossils from the Lower Oligocene of South Australia. Rec. S. 
Aust. Mus. 27(2): 117-123. 

A single, damaged cetacean tooth from the upper part of the Buccleuch Formation of the 
Murray Basin dates from the early Oligocene. It has been compared with Eocene archaeocetes 
and with late Oligocene cetaceans. Similarities are seen with Mammalodon colliveri and 
Metasqualodon harwoodii, but no definite assignment can be made. 

A limb bone from rocks of similar age in the St Vincent Basin is described briefly, but cannot 
be ascribed definitely to any particular cetacean group. 

Neville S. Pledge, South Australian Museum, Adelaide, South Australia 5000. Manuscript 
received January 1994. 



Despite its vast areas and extensive sequence of 
Tertiary sediments (e.g. Ludbrook 1969), South 
Australia seems to be remarkably deficient in 
cetacean fossils. Most of these, usually just single 
or a few associated bones of cetaceans, have been 
found in lower Miocene sediments exposed in 
cliffs along the lower part of the River Murray. 
Discoveries are summarised in Glaessner (1955), 
Pledge and Rothausen (1977), Fordyce (1982, 
1984) and Bearlin (1988). 

One of these whales, Metasqualodon harwoodii 
(Sanger 1881), was found in the Wellington area 
of the River Murray, but its exact site, and 
consequently its precise stratigraphic horizon, was 
not recorded and has been the subject of some 
speculation (Pledge & Rothausen 1977). It was 
therefore with some excitement that the author 
received a primitive- looking cetacean tooth found 
by amateur palaeontologist and collector Mr D. J. 
Barrie in February 1989 at Fred's Landing, a few 
kilometres upstream from Wellington (Fig. 1). 
Closer examination, however, has shown the tooth 
not to be Metasqualodon harwoodii, but an older, 
possibly new taxon. Further excavation at the site 
failed to yield more cetacean specimens, but 
resulted in a rich, previously unknown, 
foraminiferal assemblage being recovered 
(Lablack, pers. comm. 19/2/91, 1991 unpublished 
report). 

The specimens are registered with the South 
Australian Museum, Palaeontology Collections 
(SAM P). MUGD refers to specimens in the 
Geology Department of the University of 
Melbourne. 



Systematics 

Class Mammalia 

Order Cetacea Brisson 1762 

Suborder incertae sedis 



Genus and species indeterminate, A 

Material: 
The damaged crown of an anterior cheek tooth, 

SAMP34517. 

Locality: 

Fred's Landing, a boat launching area 3 km 
downstream from Tailem Bend, on the east bank 
of the River Murray. (Lat. 35°17*S, Long. 

139°27'E). 

Geology and Age: 

The fossiliferous beds have a very restricted 
outcrop, but have since been recognised more 
widely in subsurface sections to the southeast. 

The pale green, limonite- stained and slightly 
glauconitic, marly fine-grained limestone has a 
rich planktonic foraminiferal fauna, with key 
species Guembelitria triseriata and 
Chiloguembelina cubensis and associated species 
Sherbonina atkinsoni, Gyroidinoides sp. cf. G. 
allani, Bolivinopsis cubensis, Globigerina 
ciperoensis and Globigerina ouchitatensis 
(Lablack 1991). 

There is an invertebrate megafauna dominated 



118 



N. S. PLEDGE 




FIGURE 1 . Locality map 



by the small brachiopod Murravia catenuliformis 
and a small morph of the echinoid Scutellinoides 
sp. cf. S. patella, with occasional Waldheimia sp. 
cf. W. insolita and Magasella woodsiana 
(brachiopods) and Corystus dysasteroides 
(echinoid) fragments, Graphularia segments, 
small scallops (Chlamys), bryozoans and asteroid 
ossicles. There are also rare small shark teeth 
(Lamna sp. cf. L. apiculata and Scapanorhynchus 
maslinensis), and a broken teleost otolith. 
Unfortunately, none of these species is age 
specific, although S. maslinensis is found mostly 
in late Eocene formations (Pledge 1967). 

Lithologically, the beds are similar to the Ettrick 
Formation (middle to upper Oligocene) which 



occurs nearby. However, on the basis of the 
foraminiferal fauna, particularly G. triseriata, 
Lablack (1991 and pers. comm.) equated the unit 
in question with the lower part (Ruwarang to 
Aldinga members) of the Port Willunga Formation 
of the St Vincent Basin. 

The planktonic foraminifera are more abundant 
in the Fred's Landing section than in the Port 
Willunga local type section for the South 
Australian lower Oligocene, where the sudden 
appearance of the key species Guembelitria 
triseriata (which has a restricted stratigraphic 
range in South Australia) marks the maximum 
flooding surface T4.4 of the major transgressive 
phase following the terminal Eocene regression 



OLIGOCENE CETACEANS 



119 



(McGowran et at, 1992). The presence of G. 
triseriata and Chiloguembelina cubensis together 
restricts the age to the early Oligocene (e.g. Moss 
and McGowran 1993), and the beds to the upper 
part of the Buccleuch Formation of the Murray 
Basin. 

Description: 

Orientation of the tooth is on the basis of 
comparison with cheek teeth of Dorudon spp., 
particularly D. stromeri, and Zygorhiza kochii 
(see Kellogg 1936) where the anterior edges of 
asymmetrical teeth tend to be shorter and closer to 
vertical, and basal cingula are developed on the 
lingual faces. Insofar as that in Zygorhiza kochii 
only the upper cheek teeth have internal cingula, 
the Fred's Landing tooth may be considered to be 
an upper right anterior cheek tooth. However, 
cingula are too variable in cetaceans to be reliable 
characters and Fordyce (pers. comm. 19/8/91) has 
suggested, on the basis of its lack of lingual or 
posterior recurvature, that the tooth might be a 
lower left anterior cheek tooth, possibly equivalent 
to P 2 or P 3 . (Fig. 2) 

The tooth is not large: its preserved basal length 
is 16.3 mm, anterior width 8.2 mm, crown height 
(measured lingually) >16 mm. It is thus slightly 
larger than the figured tooth (MUGD 1874) of 




1 Omm 








4 




iiiiliiiil 



FIGURE 2. Whale tooth SAM P34517, stereopairs in: a, 
putative lingual b, labial and c, occlusal views. Scale in 
millimetres. 



FIGURE 3. Detail of occlusal view. SAM P34517. 



Mammalodon colliveri (Pritchard 1939). It is also 
quite high-crowned for its size when compared 
with other early taxa. 

The tooth is damaged posteriorly, probably 
during life since the enamel edge is rounded and 
the exposed dentine polished by tooth-on-tooth 
wear. This has removed the distal cutting edge 
and all but the uppermost denticle. All denticles 
show occlusal wear. 

The base of the crown rises sharply to its 
midpoint, corresponding to the indented groove 
that marks the union of the two roots. (The 
possibly divided distal end is not preserved.) The 
apex of the tooth is just anterior to this and is an 
acute cusp, slightly more convex labially. The 
anterior edge (carina) descends abruptly forwards 
to a small denticle about 9 mm above the anterior 
base of the enamel and continues slightly lingually 
to a smaller basal denticle (Fig. 3). From here a 
lingual cingulum curves posteriorly, parallel to the 
base of the crown, and bears a series of five tiny 
tubercles which gradually diminish towards the 
midpoint of the tooth. The tubercles reappear on 
the posterior half of the cingulum and number at 



120 



N. S. PLEDGE 



least six. The cingulum is not continuous across 
the midpoint of the face. The lingual face of the 
crown also bears a series of rather coarse, 
somewhat anastomosing ridges or wrinkles 
(cristae rugosae) in the enamel of the lower part 
— only the medial one crosses the cingulum onto 
the basal zone. The labial face has a slightly 
greater number of slightly finer ridges which 
extend higher and lower (to within 2 mm of the 
base) than on the lingual face. There is no obvious 
labial (external) cingulum. Of the posterior carina, 
only the upper-most secondary denticle is 
preserved (it is about one-third the size of the main 
cusp) together with the sharp crest joining it to the 
apex. 

Discussion: 

The antiquity of this specimen, Early Oligocene, 
makes it quite significant, since this is when it is 
considered (e.g. Barnes & Mitchell 1978, Fordyce 
1984, 1989) that mysticetes arose from the 
archaeocetes. Indeed, there are few cetacean 
specimens recorded of this age and these are listed 
by Fordyce (1992). Because of its locality, the 
tooth was initially compared to Metasqualodon 
harwoodii (specimen P8446.4) (Fig. 4b) but that 
species, although of similar size, lacks any 
indication of the cingular cusps, and it has a 
greater cristae density (Pledge & Rothausen 
1971). Preservation is also strikingly different, the 
Metasqualodon teeth being black and the new 
tooth creamy yellow in basic colour, suggesting a 
different lithologic provenance, and hence 
formation and age. 

The Fred's Landing tooth has been compared 
with descriptions and figures of those of 
archaeocetes, particularly species of Dorudon 
(Kellogg 1936, Fordyce 1985), which are much 
larger than the new specimen. Closest similarity 
in presumed overall form is seen in D. stromeri, 
e.g. M 1 in PI. 26 (3h) (Kellogg 1936) and D. osiris 
P 2 in Pi. 22 (2) {idem.). The latter figure shows the 
teeth in good detail, to indicate that the external 
face sometimes has a basal cingulum, but in any 
case does not have cingular tubercles. The greatest 
similarity of the cingulum and its tubercles is seen 
in Zygorhiza kochii, P 2 in PI. 12 (la) (idem.). 

There are some similarities with the incomplete 
premolar tooth from Waihao, New Zealand, 
referred by Fordyce (1985) to the Dorudontinae. 
This tooth (ZMT79) is considered to be a second 
premolar; unfortunately it is broken above the base 
of the enamel, so it cannot be determined whether 
a basal cingulum was present. The surface 
ornamentation of coarse, irregular cristae on the 



lingual face and somewhat finer ones on the labial 
face, together with its overall shape of a steep 
anterior edge (possibly with a basal denticle) and 
a posterior edge with at least two large denticles 
bears a similarity with the new South Australian 
specimen, which differs most noticeably in its 
smaller size and lower cristae density. 

Comparison with other published cetacean taxa 
of similar early Oligocene age has proved 
negative. There is little or no similarity with the 
possible mysticete from the lower Oligocene of 
Waikari, North Canterbury, New Zealand 
(Fordyce 1989), nor with Keyes' (1973) early 
Oligocene 'proto-squalodontid' from Oamaru, or 
Mitchell's (1989) Llanocetus denticrenatus from 
the late Eocene/early Oligocene of Seymour 
Island, Antarctic Peninsula. The later Oligocene 
cetaceans Phococetus vasconum from France, 
Platyosphys paulsonii from the Ukraine, 
Sulakocetus dagestanicus from Caucasus, 
Ke I loggia barbarus from Azerbaidzhan, 
Aetiocetus cotylaveus from Oregon, Squalodon 
?serratus from New Zealand and 'S '. 
gambierensis from South Australia (Emlong 
1966, Glaessner 1955, 1972, Mchedlidze 1984) 
likewise provide no insight into the identity of the 
Fred's Landing cetacean. This leaves 
Metasqualodon and Mammalodon to be 
considered. 

Metasqualodon harwoodii differs slightly from 
the Fred's Landing tooth (which seems 
morphologically intermediate between SAM 
P8446.1 and P8446.4) in being more symmetrical 
about the main cusp, having finer and more 
numerous cristae in the enamel, a much less 
distinct cingulum, and fewer and much smaller 
tubercles restricted towards the extremities of the 
cingulum (rather than absent only from the central 
few millimetres). Fordyce (pers. comm. 19/8/91) 
doubts that Metasqualodon is a squalodontid, and 
asks if it might not be a mysticete like 
Mammalodon. 




41 



FIGURE 4. Near-contemporary Australian cetacean 
teeth: a, Mammalodon colliveri MUGD1874, b, 
Metasqualodon harwoodii SAM P8446.4 labial (?) 
views, scale in millimetres. 



OLIGOCENE CETACEANS 



121 



Comparison with Mammalodon colliveri 
(Pritchard 1939) is frustrating, because of the 
extremely worn nature of those teeth (Fordyce 
1984, Mitchell 1989) (Fig. 4a). However, the 
isolated tooth MUGD 1874 does show 3-4 small 
cusps on that part of the lingual cingulum still 
preserved. That species differs from the Fred's 
Landing tooth in having more numerous ridges on 
the labial face of the tooth but the significance of 
this is unknown. The extreme degree of occlusal 
wear of the teeth, all in the same plane, is a 
distinctive feature, which Mitchell (1989) 
suggested might be related to a particular feeding 
strategy. In the Fred's Landing tooth, the main 
cusp is beginning to show signs of wear. Such a 
character does not seem to occur in the putative 
squalodontids Prosqualodon davidis or 
Metasqualodon harwoodii, although some 
squalondontids may show it. Fordyce (1982) has 
compared the skull of Mammalodon to that of 
Dorudon spp. and primitive mysticetes, 
concluding that while there are similarities with 
the dorudontines, Mammalodon is probably a very 
primitive mysticete. It occupies a morphologically 
intermediate position but is too recent to have been 
the ancestor of mysticetes since it is 
contemporaneous with early cetotheres (Fordyce 
1984). Mammalodon colliveri is believed to be of 
latest Oligocene age. 

On balance, the greatest similarities of the 
Fred's Landing tooth seem to be with 
Mammalodon colliveri, yet a case might be made 
that, on the few characters preserved in these 
species, it is morphologically intermediate 
between Mammalodon and Metasqualodon. The 
tooth cannot, therefore, be assigned to either taxon, 



or indeed to any particular suborder on the 
material and evidence available. 



Genus and species indeterminate, B. 

Material: 
A right radius, SAM P10875. 

Locality: 

Cliffs between Port Willunga and Aldinga Bay, 
South Australia (Fig. 1). Collected by Mr Mark 
Hagman, 1954. 

Age: 

It is unfortunate that the locality was not 
recorded more precisely, since this stretch of coast 
line encompasses the earliest Oligocene beds 
(Lindsay 1967, 1985; Lindsay & McGowran 
1986) within the Port Willunga Formation. The 
low dip of the beds (about 3 to the south) means 
that the bone could be of almost any age within 
this period. However, it had been thoroughly 
cleaned and there remained no trace of matrix that 
might have been used for micropalaeontological 
examination. On the other hand, the thoroughness 
of the cleaning suggests the bone came from a 
sandy or clayey horizon of either the Aldinga or 
Ruwarang Members of the Port Willunga 
Formation. 

Description: 

The bone, kindly identified by R. E. Fordyce (4 
February 1981) is a right radius lacking the distal 
epiphysis (Fig. 5). It is damaged along the whole 
of the posterior edge, possibly by crushing. The 




^ii|NH|iiN|iiN|ini|ini|iin|Nmiiii^ 
0mm10 20 30 40 50 60 70 80 90 100 



FIGURE 5. Cetacean right radius SAM P10875 from Port Willunga Formation, Aldinga Bay, in lateral aspect. Scale 
in millimetres. 



122 



N. S. PLEDGE 



shaft is slightly curved, convex anteriorly, and 
more convex on its outer surface. The prominent 
anterior angle seen in archaeocetes (e.g. Kellogg 
1936) has been broken off and only its roots 
remain. The inner surface of the shaft is almost 
flat at its midpoint. At the proximal end, the 
articular surface is ovate to rectangular in outline 
and slightly concave but is damaged posteriorly, 
while the distal end of the bone is lenticular in 
cross-section. The length is 190 mm overall with a 
midpoint thickness of 22 mm and diameter of 
more than 42 mm. The proximal end measures 34 
mm by an estimated 40 mm while the distal end is 
approximately 27 x 45 mm. 

A diagonal line of punctures can be seen 
midway along the outer surface of the radius, and 
can be traced in a tight arc towards the proximal 
end and back to the posterior edge. These 
punctures are interpreted as tooth marks, possibly 
from a small shark, that were inflicted before 
burial, perhaps even before death, with the attack 
on the right paddle coming from behind. 

Discussion: 

The bone shows the pachyostosis typical of 
Basilosaurus (Kellogg 1936) and sirenians. It is 
straighter and much smaller than the tibia of 
Basilosaurus cetoides (ibid.) but resembles it 
more than it does Zygorhiza kochii or Dorudon 
spp., which it slightly exceeds in size. Apart from 
this only slight curvature of the shaft, the Aldinga 
bone differs markedly from all these species in the 
lesser development of the anterior angle. 
Conversely, its curvature and development of the 
anterior angle is greater than that seen in some 
Miocene cetotheres. It is unfortunate that the 
radius is not preserved in the still-undescribed 
Mammalodon skeleton (Fordyce, pers. comm. 
19/8/91), but that would be unlikely to be of great 
help since the early odontocetes and mysticetes 
also have archaeocete-like forelimbs (ibid.). 



Conclusions 

The early Oligocene is seen to be a crucial time 
in cetacean evolution (e.g. Fordyce 1991, 1992). 
Despite it being almost impossible to establish its 
phylogenetic position, it is possible that the Fred's 
Landing tooth represents, if only morphologically, 
an intermediate stage between the Eocene 
(dorudont) archaeocetes and the succeeding 
mysticetes and odontocetes. 

The Eocene/Oligocene boundary period is also 
important palaeogeographically and palaeo- 
climatologically, with the establishment of the 
Circum-Antarctic Current (e.g. McGowran et al. 
1992; Moss & McGowran 1993; Kennett et al 
1975; Fordyce 1977, 1989b, 1992) in the early 
Oligocene. Fordyce (1992) has linked the start of 
the radiation of modern cetaceans with this event 
and its concomitant global cooling and opening of 
new feeding niches and strategies. The richness 
and abundance of the planktonic foraminiferal 
fauna of the Fred's Landing beds suggests a 
relatively open marine environment (Lablack 
1991) which is in agreement with this scenario. 

The two specimens reported here, albeit 
taxonomically and phylogenetically indeterminate, 
offer the possibility that critical specimens for 
elucidating the origins of modern whales may 
eventually be found in this part of the world. 



Acknowledgments 

I thank John Barrie for donating this specimen to the 
South Australian Museum, Karen Lablack and Brian 
McGowran for advice on foraminiferal biostratigraphy 
and Ben McHenry for brachiopod identifications. Ewan 
Fordyce made valuable comments on an early draft of 
the paper but the interpretations are mine. Debbie 
Churches typed the manuscript and Jenni Thurmer 
advised on the figures. 



References 



BARNES, L. G. & MITCHELL, E. 1978. Cetacea. Pp. 
582-602 in 'Evolution of African Mammals'. Eds. V. 
J. Maglio and H. B. S. Cooke. Harvard University 
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BEARLIN, R. K. 1988. The morphology and systematics 
of Neogene Mysticeti from Australia and New 
Zealand. (Abstract of Ph.D. Thesis). New Zealand 
Journal of Geology and Geophysics 31: 257. 

EMLONG, D. R. 1966. A new archaic cetacean from the 
Oligocene of Northwest Oregon. Bulletin of the 



Museum of Natural History, University of Oregon. 3: 
1-51 (not seen). 

FORDYCE, R. E. 1977. The development of the Circum- 
Antarctic Current and the evolution of the Mysticeti 
(Mammalia: Cetacea). Palaeogeography, Palaeo- 
climatology, Palaeoecology 21: 265-271. 

FORDYCE, R. E. 1982. A review of Australian fossil 
Cetacea. Memoirs of the National Museum of 
Victoria 43: 43-58. 

FORDYCE, R. E. 1984. Evolution and zoogeography of 



OLIGOCENE CETACEANS 



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cetaceans in Australia. Pp. 929-948 in 'Vertebrate 
zoogeography and evolution in Australasia'. Eds. M. 
Archer and G. Clayton. Hesperian, Perth, 1203 pp. 

FORDYCE, R. E. 1985. Late Eocene archaeocete whale 
(Archaeoceti: Dorudontinae) from Waihao, South 
Canterbury, New Zealand. New Zealand Journal of 
Geology and Geophysics 28: 351-357. 

FORDYCE, R. E. 1989a. Problematic Early Oligocene 
toothed whale (Cetacea, ?Mysticeti) from Waikari, 
North Canterbury, New Zealand. New Zealand 
Journal of Geology and Geophysics 32: 395-400. 

FORDYCE, R. E. 1989b. Whales, dolphins, porpoises. 
Earth Science 42(2): 20-23. 

FORDYCE, R. E. 1992. Cetacean Evolution and Eocene/ 
Oligocene Environments. Ch. 18, pp. 368-381 in 
'Eocene-Oligocene Climatic and Biotic Evolution'. 
Eds. D. R. Prothero and W. A. Berggren. (Princeton. 
U. Press). 

GLAESSNER, M. F. 1955. Pelagic fossils (Aturia, 
penguins, whales) from the Tertiary of South 

Australia. Records of the South Australian Museum 
11: 353-372. 

KELLOGG, R. 1936. A review of the Archaeoceti. 
Carnegie Institution of Washington Publication 482: 
366 pp. 

KEYES, I. W. 1973. Early Oligocene squalodont 
cetacean from Oamaru, New Zealand. New Zealand 
Journal of Marine and Freshwater Research 7(4): 
381-390.' 

LABLACK, K. L. 1991. Early Oligocene age for 
limestone from Fred's Landing, South of Tailem 
Bend, S.A.. South Australian Department of Mines 
and Energy, unpubl. Rept. Bk No. 91/56, 2 pp. 

LINDSAY, J. M. 1967. Foraminifera and stratigraphy of 
the Type Section of Port Willunga Beds, Aldinga Bay, 
South Australia. Transactions of the Royal Society of 
South Australia 91: 93-110. 

LINDSAY, J. M. 1985. Aspects of South Australian 
foraminiferal biostratigraphy, with emphasis on 
studies of Massilina and Subbotina. In 'Stratigraphy, 
Palaeontology, Malacology'. Ed. J. M. Lindsay. 
Department of Mines and Energy, Special 
Publications: 187-214. 



LINDSAY, J. M. & McGOWRAN, B. 1986. Eocene/ 
Oligocene boundary. Adelaide region, South Australia. 
Pp. 165-173 in 'Terminal Eocene Events'. Eds. Ch. 
Pomerol & I. Premoli-Silva. Elsevier, New York. 

LUDBROOK, N. H. 1969. Tertiary Period. Pp. 172-203 
in 'Handbook of South Australian Geology'. Ed. L. 
W. Parkin. Geological Survey of South Australia, 

Adelaide. 

McGOWRAN, B., MOSS, G. & BEECROFT, A. 1992. 
Late Eocene and Early Oligocene in Southern 
Australia: local neritic signals of global oceanic 
changes. Ch. 9, pp. 178-201 in 'Eocene-Oligocene 
Climatic and Biotic Evolution'. Eds. D. R. Prothero & 
W. A. Berggren. Princeton University Press. 

MCHEDLIDZE, G. A. 1984. 'General features of the 
paleobiological evolution of Cetacea'. New Delhi, 
Amerind. 136 pp. 

MITCHELL, E. D. 1989. A new cetacean from the Late 
Eocene La Meseta Formation, Seymour Island, 
Antarctic Peninsula. Canadian Journal of Fish and 
Aquatic Science 46: 2219-2235. 

MOSS, G. & McGOWRAN, B. 1993. Foraminiferal 
turnover in neritic environments: the end-Eocene and 
mid-Oligocene events in southern Australia. 
Association of Australasian Palaeontologists Memoir 
15: 407-416. 

PLEDGE, N. S. 1967. Fossil elasmobranch teeth of 
South Australia and their stratigraphic distribution. 
Transactions of the Royal Society of South Australia 
91: 135-160, 4 pis. 

PLEDGE, N. S. & ROTHAUSEN, K. 1971. 
Metasqualodon harwoodi (Sanger, 1881) — a 
redescription. Records of the South Australian 
Museum 17(17): 285-297. 

PRITCHARD, B. G. 1939. On the discovery of a fossil 
whale in the Older Tertiaries of Torquay, Victoria. 
Victorian Naturalist 55: 151-159. 

SANGER, E. B. 1881. On a molar tooth of Zeuglodon 
from the Tertiary beds on the Murray River near 
Wellington, S.A. Proceedings of the Linnean Society 
of New South Wales 5: 298-300. 



LATE QUATERNARY CHANGES IN THE MOA FAUNA (AVES ; 
DINORNITHIFORMES) ON THE WEST COAST OF THE SOUTH ISLAND, 

NEW ZEALAND 

T. K Worthy 



Summary 

The fossil moa faunas from caves on the New Zealand West Coast between Westport and 
Greymouth are described with particular reference to Madonna Cave. Site stratigraphy and 
radiocarbon dating of moa bone collagen allow dating of associated faunas. Eight moa species are 
recorded with a total of 220 individuals. Two distinct moa faunas are recognised : 1. a glacial fauna 
(10 000 - 25 000 radiocarbon years) consisting of Pachyornis elephantopus, Euryapteryx 
geranoides, and a large morph of Megalapteryx didinus. 2. a Holocene fauna (0 to 10 000 
radiocarbon years) consisting of Anomalopteryx didiformis (most common), a small morph of 
Meglapteryx didinus, Dinornis novaezealandiae, and D. struthoides. Two species, D. giganteus and 
Pachyornis australis, were very rare and are not typical components of either fauna in this area. 



LATE QUATERNARY CHANGES IN THE MOA FAUNA (AVES; DINORNITHIFORMES) 
ON THE WEST COAST OF THE SOUTH ISLAND, NEW ZEALAND. 



T. H. WORTHY 



WORTHY, T. H. 1994. Late Quaternary changes in the moa fauna (Aves; Dinornithiformes) on 
the West Coast of the South Island, New Zealand. Rec. S. Aust. Mus. 27(2): 125-134. 

The fossil moa faunas from caves on the New Zealand West Coast between Westport and 
Greymouth are described with particular reference to Madonna Cave. Site stratigraphy and 
radiocarbon dating of moa bone collagen allow dating of associated faunas. Eight moa species 
are recorded with a total of 220 individuals. Two distinct moa faunas are recognised: 1 . a glacial 
fauna (10 000 - 25 000 radiocarbon years) consisting of Pachyornis elephantopus, 
Euryapteryx geranoides, and a large morph of Megalapteryx didinus. 2. a Holocene fauna (0 
to 10 000 radiocarbon years) consisting of Anomalopteryx didiformis (most common), a small 
morph of Megalapteryx didinus, Dinornis novaezealandiae, and D. struthoides. Two species, 
D, giganteus and Pachyornis australis, were very rare and are not typical components of either 
fauna in this area. 

T. H. WORTHY, Palaeofaunal Surveys, 43 The Ridgeway, Nelson, New Zealand. Manuscript 
received 19 April 1993. 



Discrete species assemblages of moas 
associated with specific palaeoenvironments were 
recognised by Anderson (1989) and Worthy 
(1990, 1993a). Correlated with environmental 
change from the Otira Glacial to the Holocene, 
changes in the species composition of both moa 
and non-moa faunas were described by Worthy & 
Mildenhall (1989) and Worthy (1993a), from 
fossil deposits in Honeycomb Hill Cave, Oparara 
Valley, Northwest Nelson. It is reasonable to 
expect that such changes would have parallels in 
all areas of New Zealand where significant 
vegetation changes occurred between glacial and 
interglacial conditions. 

In many continental areas palaeobiogeography 
of mammals has been used to infer climate, for 
example late Pleistocene changes in Australia 
(Lundelius 1983), but only recently has the use of 
birds been advocated (Baird 1989). Baird argues 
that birds are better suited for palaeo- 
environmental reconstructions because they are 
well represented in the fossil record, can be 
identified to species level often, and identification 
of niche in modern analogues is easier. Baird 
(1991) considered the fossil bird assemblages in 
several sites to be indicative of past environmental 
conditions, and with radiocarbon dating, 
documented environmental change through the 
late Pleistocene that was consistent with patterns 
obtained by other means. 

New Zealand has a large body of data which 
indicates it had a similar pattern of vegetation 
change to Australia. These vegetation changes are 



correlated with climatic changes during the 
interglacial - glacial cycles (McGlone 1985, 
1988). As the fossil record within New Zealand 
caves is only known to extend from the Otira (last) 
Glaciation to the present, this is the period 
considered here. Palynological studies show that 
the majority of sites in the South Island were 
dominated by grassland and shrubland taxa, with 
trees less than 10% of total taxa indicating that 
forest was sparsely present, if at all, in many 
regions during the glacial maximum (22 000 - 
14 000 years ago) (McGlone 1988). The climate 
then was characterised by temperatures about 
5 °C cooler and by being considerably drier, a 
combination which effectively excluded forests. 
During the late glacial (14 000 - 10 000 years 
ago) pollen evidence suggests rapid afforestation 
occurred in many areas, at slightly different times 
depending on local conditions. For example, 
around Wellington reafforestation was much later 
than elsewhere (Lewis and Mildenhall 1985), 
perhaps a corollary of the present frequent 
exposure to southerly winds there now. In many 
South Island areas the late glacial saw grasslands 
give way to shrublands and tall scrublands. These 
changes were correlated with increased 
temperatures and increased precipitation. At the 
commencement of the Holocene (10 000 years 
ago) a return to present day temperatures and 
precipitation similar to that now resulted in rapid 
widespread vegetation change with tall podocarp 
forests becoming dominant in most lowland areas 
(McGlone 1988). 



126 



T. H. WORTHY 



New Zealand cave fossil deposits, unlike most 
swamp and all dune deposits, range in age from 
the Otira Glacial to the present so, alone, have 
potential for demonstrating faunal changes 
correlated with the above described vegetation 
changes. Demonstration of these faunal changes is 
conditional on investigation design. If a cave is 
considered to be the 'site' and all fossils 
indiscriminately associated together, or all faunas 
from caves within a region considered as one, then 
by investigation design, perception of temporal 
changes is ruled out. Dating is necessary to 
determine the time span of deposition for each 
discrete site in a cave. In this regard isolated 
skeletons, or bones in surface deposits within a 
cave, should be considered as discrete events 
which need to be individually dated, unless on the 
basis of sedimentary history within the cave, or 
similar speleogenetic evidence, an upper age limit 
can be set on fossil deposition. 

The following example illustrates how study 
design compromises resultant conclusions. 
Atkinson & Millener (1991) analysed the cave 
fossil fauna around Waitomo in the North Island 
using data compiled from sites in 156 separate 
caves (Millener 1981), not 37 as they stated. 
Insufficient attention was paid to the age of the 
fossils. Most were collected by amateurs from 
surface deposits that could have been up to 
100 000 years old (speleothems of this age are 
known in some Waitomo caves (Hendy 1969)). 
Younger fossils were more likely to have been 
more abundant for two reasons. Firstly, younger 
fossils would have been subject to destructive 
processes for a shorter period of time, and 
secondly in many cases younger sediments overlie 
and therefore obscure older material. No attempt 
was made by these authors to justify temporal 
association of fossils from individual sites within 
caves - caves were treated as sites - nor was 
temporal association of deposits from separate 
caves proven, beyond the observation that dates 
for fossils from 10 caves ranged from one to 
twenty five thousand years old. Despite the proven 
age of the fossil deposits spanning two major 
climate regimes, all species in the faunal list were 
assumed to have been present contemporaneously 
between 6 000 and 1 000 years BP. To treat all 
fossils as a unit necessarily obscured any patterns 
there may have been and so, as younger fossils 
predominated, Atkinson and Millener' s approach 
resulted in the conclusion: 'A humid lowland 
forest thus dominated the area throughout the 
period when faunal remains were accumulating'. 
This is a description of the environment that 



prevailed in the area during the Holocene, and 
describes conditions that must have differed from 
those prevailing during the glacial period, if the 
palaeotemperature curve for the region is any 
indication (Hendy 1969). 

Apart from age, the limitations imposed by 
taphonomic processes (Baird 1991), especially as 
they apply to species representation, need to be 
understood and assessed before sites are 
compared. With increased water flow at a site 
there is progressive removal and destruction from 
smaller to larger fossils resulting in a marked bias 
towards preservation of moa bones in stream 
sediments. The New Zealand birds observed to be 
specific indicator species of open-country habitat, 
for example the pipit and New Zealand quail, have 
small bones for which suitable preservation 
conditions are rarely encountered in New Zealand 
caves. Their absence or rarity in the Waitomo 
fauna as a whole (as so far described) does not 
preclude their having been common in the area 
during the Otiran. 

The present study arose out of a survey of the 
Quaternary fossil bird fauna of the Punakaiki karst 
region, West Coast, South Island (Worthy & 
Holdaway 1993). Preliminary examination of the 
Canterbury Museum's collection of fossils from 
the region indicated that elements of both the 
Anomalopteryx and Euryapteryx assemblages of 



NORTHl AND 



HONEYCOMB HILL CAVE 




FIGURE 1. Locality map showing the study area in New 
Zealand. 



128 



T. H. WORTHY 



Fossil Deposits 



Results transported fossils that were not enclosed in 

sediments (sites 1-9,14,15); 2. Allochthonous 

deposits of fossils that were in fluvial sediments 

(sites 11, 11a, 13, 16); 3. Fossils that had 

accumulated in autochthonous deposits below 

The fossil sites located within Madonna Cave entrance shafts or lay, as individual skeletons, near 

are shown on Figure 2. There were three main an entrance. Limited water transport (2-10 m) of 

types of fossil deposits in the cave: 1. some fossils had occurred at some sites, for 

Allochthonous deposits of scattered water- example The Morgue, (sites 10, 12, 13a, 17, 18, 19). 



MADONNA - EQUINOX CAVE SYSTEM 
WAGGON CREEK - TIROPAHI RIVER 



!5 - 16 



Nmag 




tNl HANCF 



EQUINOX CAVE 

Length 2S0m 



FNTRANCL 



MADONNA CAVE 

Length 1436m 




FIGURE 2. Map of the Madonna: Equinox Cave System showing the location of fossil sites 1-19 referred to in the 
text. 



CHANGES IN MOA FAUNAS 



127 



Anderson (1989) occurred in apparent geographic 
sympatry which would have been inconsistent 
with observations made by Worthy (1990). Further 
north at Honeycomb Hill Cave on the West Coast 
recent research has shown that moa assemblages 
changed with climate from the Otiran to the 
Holocene (Worthy & Mildenhall 1989; Worthy 
1993a). I therefore thought it probable that the 
fossil deposits of the Punakaiki area also preserved 
elements of at least two successive and distinct 
moa faunas. 

During the survey of caves in the Punakaiki area 
a site was searched for in which to test the 
hypothesis of moa faunal turn-over. I chose 
Madonna Cave because it contained several 
closely associated sites, each with stratigraphic 
control, and large numbers of fossils. The initial 
examination of the cave revealed a number of rich 
and discrete fossil sites representing a range of 
depositional environments. Some were in fluvial 
stratified sediments, and others under entrances. 
Different species assemblages were noted in 
different sites. In addition, it was noted that 
discrete periods of sedimentation had incorporated 
fossils in stratified sediments that were therefore 
likely to provide dates reliable for building a 
chronology of change in moa species. Later 
erosion had re-excavated most of these sediments 
leaving only remnant banks, and because these 
fluvial sediments had also been deposited and 
eroded from passages beneath entrance shafts, the 
fossil deposits under the shafts must have 
accumulated subsequent to the erosion. Therefore, 
data from these various sites could be used, in 
combination, to test the hypothesis. 

Madonna Cave is between Waggon Creek and 
Doubtful Creek on the south side of the Tiropahi 
(Four Mile) River, between Westport and 
Greymouth (Fig. 1). The approximate grid 
reference for the centre of the cave system, on the 
metric 1: 50 000 map series NZMS 260, is K30 
819127. The cave system drains a karst area, 
about 200 m above sea level, that has many 
dolines and natural traps in the form of shafts and 
grikes. The area is vegetated in mature mixed 
beech/podocarp forest with emergent rata trees up 
to 35 m in height. The forest floor and tree trunks 
are cloaked in deep moss that is indicative of the 
regular high rainfall (c. 3 000mm). 

The cave system comprises three sections: 
Equinox Cave about 250 m long; and the 'upper 
levels' and the 'lower levels* of Madonna Cave 
which together have 1 436 m of surveyed passages 
in them. The cave is characterised by numerous 
entrances (see Fig. 2). In each area of the cave 



small streams now flow in the lowest levels but it 
is in the older stream courses at higher levels that 
sedimentary deposits are preserved. These 
sediments vary from sands to coarse gravel and 
moa fossils are visible in some. Abundant moa 
fossils also occurred in the streambeds. 



Definitions 

Allochthonous versus autochthonous. Fossils in 
caves have been classified as allochthonous (died 
at a different place to that in which the fossils are 
deposited), or autochthonous (where the fossils are 
found at or near where the animals died) by Baird 
(1991). In the surface environment such an 
assessment of fossil sites prior to 
palaeoenvironmental reconstruction is extremely 
important as fossils in fluvial deposition sites 
could have their origin many miles away, and 
therefore could have been transported out of the 
environment they lived in. Assessing cave sites in 
this fashion would result in fossils from stream 
sediments being classed as allochthonous. 
However, I contend that to discount use of data 
from these sources for palaeoenvironmental 
reconstruction, as is necessary for surface sites, 
would omit relevant data. This view is held as in 
the majority of caves, and especially within 
Madonna Cave, all fossiliferous fluvial sediments 
can be easily traced to their point of origin that is 
at most only a few hundred metres distant. So all 
bird fossils in such cave sediments can be 
considered autochthonous with respect to the 
home range of the living bird, and can justifiably 
be used in palaeoenvironmental reconstructions. I 
therefore use the terms 'allochthonous deposits' 
and 'autochthonous deposits' to indicate only that 
bones have, or have not, been secondarily 
transported from the point of death. I use the term 
'autochthonous assemblage' to indicate that it is 
an assemblage of local derivation, which may be 
in either of the above deposit types. 

MNI is minimum number of individuals 
determined from the number of the most abundant 
ipsolateral element. 

Radiocarbon dating was done by the Institute of 
Geological and Nuclear Sciences Ltd laboratories 
at Lower Hutt, New Zealand, on the collagen 
fraction of bones, and ages are given as 
'conventional radiocarbon age', where Libby TVi 
= 5568 yrs BP. 

MNZ is the Museum of New Zealand Te Papa 
Tongarewa (was the National Museum of NZ) 
Wellington. 



CHANGES IN MOA FAUNAS 



129 



1. Water transported fossils. 

Main Stream. Fossils at sites 1-8 were found in 
allochthonous deposits as isolated bones in the 
floor of the streambed. After recording all fossils 
it was decided that generally only femora would 
be collected leaving other fossils in situ for future 
cave visitors to see. The fossils collected 
represented all species and the maximum number 
of individuals present in sites 1-8 (MNZ S28055- 
28065) (Table 1). The exception to the policy of 
collecting femora was in site 8 where a 
tarsometatarsus of Pachyornis (S28064) was 
collected since this tax on was otherwise 
unrepresented. The majority of fossils were little 
worn and were whole bones, but at the most 
downstream part of the stream (site 8) a few very 
worn fragmentary bones of species not otherwise 
represented in the fauna were found. These 
included a distal left femur and a distal left 
tibiotarsus of Pachyornis and a distal right 
tarsometatarsus of E. geranoides. 

The age of the majority of fossils in sites 1-8 
was considered likely to be Holocene with the few 
worn specimens (detailed above) interpreted as 
probably reworked from older sediments. One 
bone of A. didiformis K30/f62b (assumed 
'young'), and another, part of P. elephantopus 
S28064, K30/f62a, (assumed 'old') were dated to 
test this (Table 2). 

Near site 1 a small side-passage on the true 
right had several fossil moa bones scattered along 
its terminal crawlway. These were not collected 
and were considered to be a sub-sample of the 
fauna in the main stream sites 1-8. Bones of 



Dinornis novaezealandiae and Anomalopteryx 
didiformis were seen. 

Fossils at sites 14 and 15 lay in the bed of the 
stream. Both sites were similar in that they were 
about 10 m above the Main Stream in passages 
whose sediment banks had a different sequence of 
sediments to that seen in the Main Stream. I 
interpreted this to mean that the passage around 
sites 14 and 15 had escaped the last period of 
sedimentary infill and re-excavation that the Main 
Stream had been subjected to. As the small 
streams in these passages come from shaft 
entrances it was postulated that moas of both 
Holocene and pre-Holocene derivation could be 
present. All specifically recognisable bones were 
collected from site 14 and a Pachyornis 
elephantopus bone, chosen because this taxon was 
not present in the assumed young sites 10 and 12, 
was dated. Fossils found in the stream bed at site 
15 were left as reference specimens. 



2. Fossils enclosed in fluvial sediments 

All along the Main Stream there were remnant 
banks of sediment reflecting an earlier period of 
sedimentation followed by erosion. Near site 8 two 
bones were observed apparently in situ near the 
top of the sediment bank: one an Anomalopteryx 
didiformis femur was left in situ, and the other, a 
femur of a juvenile Dinornis MNZ S28218, was 
collected. Opposite where the stream sinks another 
fossil (MNZ S28079), of undetermined moa 
species, was collected from about 0.5 m below the 
top of the sediments. Because these sediments 



TABLE 1. The identity of 63 moas represented in collections from Madonna Cave, or by in situ material, and their 
distribution within sites is summarised here. Specimens marked * represent individuals known from in situ material 
only. All the rest are represented by collected specimens that were chosen to represent the total material from the site 
and so MNI at each site would not increase with further collecting even though many bones remain. 



Site 


A. did. 


M. did. 


D. str. 


D. no v. 


D gig- 


E. ger. 


P. ele. 


1-8 


6 





2 


2 





1* 


1 


9 





3 


1 














10 


5 


5 


2 


8 


1 








11 





1 














3 


12 


3 


1 


1 


1* 











13 





1 


f) 








1 


1 


13a 


1 




















14 


3 





I 











1 


15 











1* 








1* 


16 

















2 


1 


17 











1* 











19 


1* 




















Total 


19 


11 


7 


13 


1 


4 


8 



130 



T. H. WORTHY 



border an active stream and are no more than 
0.5 m above the normal water level they are 
probably considerably younger than those 
hereafter described. 

Site 11. This site was at the south end of 
Material World. The limestone floor of the passage 
was visible at the end of this passage but remnant 
sediment banks extended to the roof on both sides. 
At site 1 1 the sedimentary sequence was as 
follows: about 0.5 m of stream gravels and sands 
rested on the limestone floor. The gravels were 
overlain by a 0.5 m thick layer of slightly rounded 
limestone cobbles embedded in sand which was, 
in turn, overlain by homogenous sands that filled 
the 0.2-0.3 m space between the cobble layer and 
the roof. Immediately west of site 1 1, a 2 m bank 
overhung a streamway, but the passage leading 
upstream via the crawl (see map), preserved the 
sequence of sediments very well. Also, half way 
up the passage towards Circuit Tomo, an alcove 
preserved the same sedimentary sequence. There 
the available roof space above the cobbles was 
much greater and the sand layer much thicker. 
Immediately east of The Morgue the passage roof 
was also much higher and the sedimentary 
sequence was preserved in several places showing 
the sand layer was at least 3 m thick. Fossils were 
found at the interface of the sand and cobble layers 
: two bones (MNZ S28076, S28080) at site 11, 
and one at site 11a that was left in situ. Two 
fossils (MNZ S28077-8) were recovered from the 
crawl immediately west of site 1 1. Although not 
in situ preservation characteristics and adherent 
matrix indicated they had been dislodged from the 
same layer. Part of the M. didinus femur collected 
from in situ in site 1 1 was dated. 

Site 13. Stream laid sand and gravel banks 



indicated that this area of the cave had been 
infilled to the roof. Worn fossil bones from single 
individuals of Megalapteryx MNZ S28082, 
Pachyornis MNZ S28081 and Euryapteryx MNZ 
S28083 were scattered on the surface of the floor 
formed on the eroded surface of these sediments. 
In one of the remnant banks of sediment lining the 
site, in situ fossils from these individuals were 
recovered, indicating that those fossils on the 
surface had been eroded from the sediments. One 
of the eroded Euryapteryx bones recovered from 
the surface of this site was dated. 

Site 16. Here exactly the same sequence of 
sediments as seen at site 1 1 was preserved. Some 
fossils (MNZ S28121) were exposed on the 
surface but excavation revealed that other fossils 
were deposited on top of the limestone cobble 
layer (MNZ S28220-223). Several were found 
embedded in sand adjacent the upstream end of a 
large fallen rock where the stream had apparently 
washed them. About one metre 'upstream' of this 
rock, a remnant bank of sediment overlying the 
cobbles was found to contain an articulated series 
of moa vertebrae MNZ S28224 at a depth of three 
to four centimetres and about 30 cm above the 
sand cobble interface. These are most probably the 
surviving parts of the skeleton otherwise 
represented by S28121. This skeleton was 
deposited near the end of the deposition period as 
a whole carcass. The femur of S28121 was dated 
to provide a minimum age for the site and the 
more deeply buried disarticulated material. 



3. Fossils accumulated near entrances 

The most significant site was The Morgue, site 



TABLE 2. Radiocarbon dating results from Madonna Cave. Specimens are identified by species, site of origin, Fossil 
Record Number FRN, and the laboratory sample number NZA (Accelerator mass spectrometry laboratory) or NZ 
(Gas counting laboratory). 



Site 


Species 


FRN 


NZAor 

NZ 1 


Radiocarbon 

age 


8 


P. elephantopus 


K30/f62a 


2505 


14740±110 


8 


A. didiformis 


K30/f62b 


2443 


2197±86 


10 


A. didiformis 


K30/f65a 


2506 


5447±87 


10 


D. novaezealandiae 


K30/f65b 


7925 1 


5893±88 


10 


D. giganteus 


K30/f65c 


7926 1 


2829±75 


10 


M. didinus 


K30/f65d 


2503 


782±83 


11 


M. didinus 


K30/f67 


2780 


I3150±140 


12 


A. didiformis 


K30/f66 


2521 


1076±83 


13 


E. geranoides 


K30/f63 


2779 


1 1090+100 


14 


P. elephantopus 


K30/f64 


2446 


20680±160 


16 


E. geranoides 


K30/f61 


2445 


23780±210 



CHANGES IN MOA FAUNAS 



131 



10. Here an entrance shaft at least 15 m deep was 
the natural trap that had accumulated a large 
amount of material in an autochthonous deposit. 
The history of sedimentation in the adjacent 
passages was one of deposition, wherein the 
sequence recorded for site 1 1 was deposited, 
followed by erosion right back to floor level to 
leave only sediment remnants along some walls 
and in alcoves. Only following this erosion event 
could the fossils observed at The Morgue have 
started to accumulate, and therefore they must all 
be younger than those seen in site 1 1. The base of 
the entrance shaft intersects a passage aligned 
approximately north-south which, either side, is 
only 0.5 m high. The water that formed the shaft 
now drains via a 20-30 cm high bedding-plane 
passage to the east. Although impenetrable this 
bedding can be approached via a small passage 
from the east that starts at the top of a 5 m drop, 
below which the water ultimately sinks away in 
gravels. Fossils were mostly collected from the 
base of the entrance shaft and from the bedding 
plane passage and its downstream outlet (MNZ 
S28088-28112, 28114-28120). Articulated 
remains of one subadult Dinornis 
novaezealandiae and one Anomalopteryx 
didiformis were found in the passage to the south 
indicating that these trapped birds had passed the 
low passage in this direction. Immediately north 
of the base of the entrance shaft an articulated 
skeleton of A. didiformis was left in situ. Fossils 
found at site 9 scattered along the floor of Material 
World are interpreted as remains of birds that were 
trapped by the Morgue entrance and passed the 
low passage to the north. It is significant that these 
are mainly from 3 individuals of Megalapteryx, 
the smallest moas in the region (MNZ S28067- 
28071). These small M. didinus, like others in site 
10, are a small morph of the species identified by 
Worthy & Holdaway (1993) as only present in 
Holocene deposits in the area. One specimen in 
site 9 was a very worn, possibly reworked, femur 
of Dinornis (MNZ S28066). Four bones were 
dated from site 10, one from each of the following 
taxa: Megalapteryx didinus, Anomalopteryx 
didiformis, Dinornis novaezealandiae, and D. 
giganteus (from S281 14). 

Circuit Tomo: site 12. This site was at the base 
of another shaft entrance. Fossils were located in 
an autochthonous deposit of rockfall debris at the 
shaft base or scattered down the first few metres 
of the stream passage away from the shaft. 
Sediment banks with the same stratification as 
sediments preserved at site 1 1 indicate that fossils 
in site 12 had the same potential age range as 



those in site 10, and had to be younger than those 
in site 11. The M. didinus specimen was one of 
the small morphs of the species. 

Other sites. A few fossils were recovered from 
site 13a. Bones of an individual A. didiformis 
were left in situ near the upstream entrance to the 
Main Stream (site 19). In Equinox Cave at site 17 
a complete D. novaezealandiae skeleton was left 
in situ lying on the surface, and is necessarily 
younger than the fossils enclosed in sediments 
nearby at site 16. Closer to the entrance a small 
crawlway leads off and it contained some fossils, 
but none of moas. 



Summary of Moa Distribution in Madonna 
Cave 

Table 1 lists the individual sites described 
above and the moas, by species, from each of 
them: 63 individual moa are represented in the 
total collection. The geological ages for each site 
are indicated from the dates listed in Table 2. Sites 
11, 14 and 16 are exclusively Otiran in age and 
contain Pachyornis, Euryapteryx, and 
Megalapteryx. Sites 10 and 12 are exclusively 
Holocene, and have Anomalopteryx, three 
Dinornis species with D. novaezealandiae the 
most abundant, and Megalapteryx. From the 
cave's sedimentary history site 9 should be 
associated with 10 and 12, and so is of Holocene 
age. Three dates were obtained from fossils found 
loose in streambeds; one from site 15, and two 
from the Main Stream sites 1-8. In the Main 
Stream the Anomalopteryx was late Holocene and 
in the Pachyornis Otiran. In site 15 the 
Pachyornis bone that was dated was Otiran in 
age. 



Discussion 

Allochthonous and autochthonous deposits in 
Madonna Cave contain fossils of various ages that 
can be regarded as autochthonous assemblages, 
that because of known age ranges provide useful 
palaeoenvironmental indicators. Deposits beneath 
shafts from the surface contain only Holocene 
faunas, fluvial sediments not in active streamways 
contain Otiran faunas, and streamways contain 
mixed Otiran and Holocene faunas. Holocene 
faunas were characterised by abundant 
Anomalopteryx didiformis, Dinornis 

novaezealandiae, D. struthoides, and small to 
medium morphs of Megalapteryx didinus. The 



132 



T. H. WORTHY 



Otiran fauna contained no A. didiformis, but 
common Pachyornis elephantopus and 
Euryapteryx geranoides, with M. didinus as a 
medium to large morph only. 



fewer surviving Otiran sites. P. australis and D. 
giganteus were very rare, and so in the Punakaiki 
karst were considered to have been outside of their 
preferred geographic ranges. 



Local comparisons 

A series of dates was obtained from moa bones 
from Te Ana Titi (Worthy & Holdaway 1993). 
The single Pachyornis australis individual was 
Otiran in age, and the Dinornis novaezealandiae 
of Holocene age. The 18 Megalapteryx didinus 
recorded from Te Ana Titi ranged from the largest 
ever recorded to nearly the smallest. The dates 
obtained by Worthy & Holdaway (1993) clearly 
showed that all the largest, without exception, 
were of Otiran age and the smallest, of Holocene 
age. These data support the hypothesis of post- 
glacial dwarfing proposed by Worthy (1988). It is 
therefore significant to note that the Megalapteryx 
recorded from the Holocene sites 9, 10 and 12 in 
Madonna were all small individuals. 

Table 3 lists all moa individuals recorded from 
the Punakaiki karst area by Worthy & Holdaway 
(1993) regardless of specimen age. A. didiformis 
was the most abundant, with nearly twice the 
number of the next most frequently recorded 
species. This reflects the preponderance of 
Holocene sites and that this species was the most 
common in them. Megalapteryx didinus, the only 
species occurring in both Otiran and Holocene 
deposits, was the next most frequently 
encountered. Dinornis struthoides and D. 
novae zealandiae, the remaining common 
Holocene species were the next most abundant. 
The Otiran Pachyornis and Euryapteryx occurred 
in lower frequencies consistent with there being 



Comparison with other areas 

The only place where comparable data has been 
obtained is Honeycomb Hill Cave System in the 
Oparara Valley, northwest Nelson. This area 
ranges in altitude from 200 - 300 m a.s.L There 
the Otiran moa fauna was dominated by 
Pachyornis elephantopus, P. australis and 
Megalapteryx didinus. Euryapteryx geranoides 
was extremely rare (one individual among 
hundreds), and Dinornis novaezealandiae and D. 
struthoides were both rare. In the Holocene 
Pachyornis and Euryapteryx were not recorded 
and Anomalopteryx didiformis dominated with D. 
novaezealandiae becoming more common 
(Worthy & Mildenhall 1989, Worthy 1993a). Data 
documenting changes in moa fauna over time have 
not been published for any other area. 

Worthy (1990) described the 'Holocene' moa 
faunas for many regions in New Zealand. The 
Holocene faunas of western South Island, as 
shown by Honeycomb Hill Cave and the 
Punakaiki karst, were similar to those in lowland 
wet forest areas in Southland, and central, western 
areas of the North Island with Anomalopteryx 
didiformis and Dinornis novaezealandiae 
dominating in both. The Otiran faunas of the 
western South Island were most like the Holocene 
faunas of eastern areas of the South Island but 
there were significant differences. As seen above, 
in the west, Megalapteryx didinus accompanied 
Pachyornis elephantopus and Euryapteryx 



TABLE 3. Summary of total numbers of moa individuals recorded from the Punakaiki karst area by Worthy & 
Holdaway (1993). MNI in coll. is minimum number of individuals determined from specimens in museum collections 
(some of which are voucher specimens for more material in the fossil site). MNI in situ refers to those recorded from 
the cave site only, and for which no voucher specimen was collected. 



Dinornithiformes 


MNI in coll. 


MNI in situ 


Total No. 


(%) 


Anomalopteryx didiformis 


75 


6 


81 


(40.3) 


Megalapteryx didinus 


28 


10 


38 


(18.9) 


Pachyornis elephantopus 


11 




11 


(5.5) 


Pachyornis australis 


3 




3 


(1-5) 


Euryapteryx geranoides 


17 




17 


(8.4) 


Dinornis struthoides 


24 


I 


25 


(12.4) 


Dinornis novaezealandiae 


22 


2 


24 


(11.9) 


Dinornis giganteus 


2 




2 


(1-0) 


TOTAL 


182 


19 


201 





CHANGES IN MOA FAUNAS 



133 



geranoides, but in the east, although the latter two 
species were common, they were accompanied by 
Emeus crassus (unknown in the west) and D. 
giganteus (very rare in the west). 

The western Otiran fauna was more similar to 
that in Otiran loess deposits in eastern South 
Island where Pachyornis elephantopus dominated 
with Euryapteryx geranoides the next most 
abundant species. A. didiformis, M. didinus, and 
D. novaezealandiae were not present and E. 
crassus, D. giganteus and D. struthoides were 
rare (Worthy 1993b). On Takaka Hill in northwest 
Nelson the Otiran fauna consisted of rare M. 
didinus, and common Pachyornis elephantopus, 
P. australis and E. geranoides (Worthy unpubl. 
data). 

The western distribution of moas in their dated 
context support previous notions of habitat 
preference of individual species (Worthy 1990). 
The mainly western distribution of M. didinus in 
both Otiran and Holocene periods suggests that it 
preferred wetter and cooler habitats. Generally it 
was an upland bird of the montane forests to the 
subalpine zone (Worthy 1988, 1989b). However, 
in the Holocene forests of the Punakaiki area, the 
only known lowland population of M. didinus was 
represented by a race where individuals were 
smaller than Anomalopteryx and their upland 
counterparts. It is possible that the small size of 
individuals enabled this population to avoid 
competition with A. didiformis which was the only 
other emeid present. Elsewhere populations of 
these two species met in ecotones in the montane 
forest zone and were separated by altitude (Worthy 
1990). P. australis was rare in both time periods 
in Punakaiki. An abundance in Otiran Honeycomb 
Hill Cave and Takaka deposits compared to its 
presence mainly in subalpine deposits during the 
Holocene indicate that it was an upland species 
(Worthy 1989a, 1989b). In contrast P. 
elephantopus and E. geranoides were both 
common during the Otiran in a range of areas, 
including lowland eastern areas and western sites 
from sea-level to 700 m, but during the Holocene 
were only in eastern dry lowland areas. These 



observations support the hypothesis that these taxa 
preferred relatively openly vegetated areas where 
mosaics of grass and shrubland dominated the 
vegetation. 



Conclusion 

The data presented here clearly show that there 
were marked changes in distribution of moa 
species on the West Coast of the South Island 
between the last Glacial period and the Holocene. 
Usually only 3 to 4 species co-existed at one time 
and so moa species assemblages could be used as 
stratigraphic indicators. The presence of E. 
geranoides and P. elephantopus are indicative of 
Otiran age deposits and A. didiformis and D. 
novaezealandiae Holocene deposits. In addition 
to changes in species distribution M. didinus 
exhibited marked post-glacial dwarfing. The 
smallest and largest forms (defined Figure 26, 
Worthy & Holdaway 1993) are therefore useful 
stratigraphic indicators for the Holocene and 
Otiran periods respectively. The national 
implication of these data is that faunas from 
different sites should not be amalgamated into a 
regional fauna and analysed as a unit without 
reference to temporal variation unless temporal 
association is first proven. Cave systems 
throughout New Zealand can be expected to have 
superimposed Otiran and Holocene faunas and 
only careful study of stratigraphy combined with 
dating will separate them. 



Acknowledgments 

The study was funded by a grant from the Foundation 
for Research, Science and Technology. Dating of bone 
samples was supported by a grant from the Lottery 
Science Research fund administered by the NZ Lottery 
Grants Board. It would not have been possible without 
the hard work of my field assistant Anne Melhuish, nor 
the support of the Department of Conservation on whose 
land the study site lies. 



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AN EXTINCT SPECIES OF CORMORANT (PHALACROCORACIDAE, 
AVES) FROM A WESTERN AUSTRALIAN PEAT SWAMP 

G. F. Van Tets 



Summary 

Microcarbo serventyorum n.sp. is described from a pelvis with proximal parts of the femora and 
some caudal vertebrae, which were found in a peat swamp at West Bullsbrook, north of Perth, 
Western Australia. Age is not known, but is probably Holocene. These bone exaggerated the pelvic 
differences between those of small extant species of Phalacrocorax and Microcarbo and are 
therefore included as a new species in the latter genus. Presumably M. seventyorum was able to 
forage in even more confined waters than other species of Microcarbo. 



AN EXTINCT NEW SPECIES OF CORMORANT (PHALACROCORACIDAE, AVES) 
FROM A WESTERN AUSTRALIAN PEAT SWAMP 

G. F. VAN TETS 

van TETS, G. F. 1994. An extinct new species of cormorant (Phalacrocoracidae, Aves) from a 
Western Australian peat swamp. Rec. S. Aust. Mus. 27(2): 135-138. 

Microcarbo serventyorum n.sp. is described from a pelvis with proximal parts of the femora 
and some caudal vertebrae, which were found in a peat swamp at West Bullsbrook, north of 
Perth, Western Australia. Age is not known, but is probably Holocene. These bone exaggerated 
the pelvic differences between those of small extant species of Phalacrocorax and Microcarbo 
and are therefore included as a new species in the latter genus. Presumably M. serventyorum 
was able to forage in even more confined waters than other species of Microcarbo. 

G. F. van Tets, Wildlife & Ecology, CSIRO, P.O. Box 84, Lyneham, Australian Capital 
Territory 2602 and Division of Archaeology and Natural History, Research School of Pacific 
and Asian Studies, Australian National University, Australian Capital Territory 0200. 
Manuscript received 13 April 1993. 



At the 16th International Ornithological 
Congress in Canberra, I postulated that 
Australasia might be the main centre of generic 
and sub-generic divergence of the extant 
Phalacrocoracidae (van Tets 1976). Since then 
Siegel-Causey (1988) raised my generic separation 
of cormorants, Phalacrocorax versus shags, 
Leucocarbo to that of the subfamilies 
Phalacrocoracinae and Leucocarbinae. I agree 
with these subfamilies (van Tets pp. 808-809 in 
Marchant & Higgins 1990), but disagree with 
some of Siegel-Causey' s generic and specific 
arrangements in those subfamilies. In particular I 
accept as cormorants in the Phalacrocoracinae only 
Microcarbo and Phalacrocorax including the 
sub-genus Hypoleucos. 

Evidence for a new extinct Australian species of 
cormorant is provided by a pelvis with proximal 
parts of the femora and some caudal vertebrae 
(Figs 1-4). They were found 9 January 1970 in a 
peat swamp at mining lease 19H 'Melaleuca', 
West Bullsbrook, at 30 km north of Perth, 
Western Australia (31°41'S 115°59'E). They were 
noticed on a stockpile, before loading on to a 
truck, but after rotation and blading, from an 
estimated depth of about one foot (=0.3 m). Age is 
not known, but is probably late Holocene. C. 
Mizen forwarded it per A. R. Burns to Duncan 
Merrilees at the Western Australian Museum, who 
registered it 12 February 1970 as WAM 70.2.10. 

WAM 70.2.10 was compared with a wide range 
of material in the collections of the Australian 
Museum (AM) Sydney, the Australian National 
Wildlife Collection (ANWC) CSIRO Canberra 
and the University of Michigan Museum of 



Zoology (UMMZ) Ann Arbor. The material 
included all families and genera of Pelecaniformes 
and most species of Phalacrocoracidae including 
all of Microcarbo (sensu van Tets 1976). 



System atics 

In penguins (Spheniscidae), darters 
(Anhingidae), cormorants and shags 
(Phalacrocoracidae), the cranial facet of the pelvis 
is convex and not concave as in frigatebirds 
(Fregatidae), gannets and boobies (Sulidae), nor 
saddle-shaped as in pelicans (Pelecanidae), 
tropicbirds (Phaethontidae) and in almost all other 
kinds of birds. 

The pelves of the Spheniscidae differ from those 
of the Anhingidae and Phalacrocoracidae by 
having a fenestrated preacetabular ala of the ilium 
and lacking ventral spines (van Tets & O'Connor 
1983). On the pelves of the Anhingidae prominent 
ilial crests flank the median dorsal ridge and 
shield (Olson 1975) and on those of the 
Phalacrocoracidae only the caudal half of the 
shield. In the shield there are up to six pairs of 
postacetabular foramina in Anhingidae and up to 
eight in Phalacrocoracidae. These foramina are 
relatively smaller in Anhingidae than in 
Phalacrocoracidae. 

On the femur, the trochanter is about as tall as 
the head in Phalacrocoracidae and not lower as in 
WAM 70.2.10, Anhingidae and Sulidae (van Tets 
etal. 1989). 

On the ilia lateral to the first pair of 
postacetabular foramina is a pair of rough 



136 



G. F. VAN TETS 



; 




srlf T 





MM I" ij"Mi]iii[iiiiMiiijiiiiiiin||iiHiiii] 

FIGURE 1. Dorsal view of pelvis. Left: Phalacrocorax 
sulcirostris ANWC BS-1310. Right: Microcarbo 
melanoleucos ANWC BS-1419. 



irregular 'facets', which are about as wide as 
broad in Phalacrocoracinae and are longer than 
broad extending alongside the second pair of 
foramina in Leucocarbinae, but are variable and 
intermediate in the Pied cormorant, 
Phalacrocorax varius, and the Black-faced shag, 
Leucoca rbo fusee scens. 

The first four pairs of postacetabular foramina 
are about twice as long as wide in Phalacrocorax 
and relatively broader and less than twice as long 
as wide in Microcarbo. The concave preacetabular 
lateral edge of ilium is relatively long in 
Microcarbo and short in Phalacrocorax. The 
pelvis is relatively longer and more slender in 
Phalacrocorax than in Microcarbo. 

Because WAM 70.2.10 fits the above characters 
of Microcarbo, except for the femora, I propose a 
new specific name for it. 

Microcarbo serventyorum sp. nov. 

Etymology 

In honour of the Serventy brothers, Dom and 
Vincent, for their contributions to our knowledge 




mm 



-■v- 




MM 



miiiiii i imiim i iiiiiiMiiiiiinmmiiiiTTT 



FIGURE 2. Dorsal view of pelvis. Microcarbo 
serventyorum WAM 70.2.10. 



of the Australian cormorants and shags (Serventy 
1938, 1939, Serventy & White 1943, Serventy et 
al. 1971). 

Diagnosis 

A cormorant slightly smaller than species of 
Microcarbo and Phalacrocorax, relatively long 
preacetabular concave lateral edge on ilium; 
relatively broader first four pairs of postacetabular 
foramina adjacent to narrower ilia, and much 
larger 8th pair than in species of Microcarbo and 
Phalacrocorax. Parapophyses between 7th and 
8th pair extend cranio-laterally and not laterally as 
in Microcarbo and Phalacrocorax (See figs 1—4). 

Measurements of WAM 70.2.10 are generally 
more than ± 3 sd away from mean measurements 
of the smallest extant species of Phalacrocorax, 
and differ from all extant species of Microcarbo. 
These differences are most marked in the length of 
the dorsal ridge of the pelvis, the length of the 8th 
postacetabular foramen, and the proximal width of 
the femur (Table 1). 



EXTINCT NEW CORMORANT 



137 





FIGURE 3. Caudal view of left femur. Left: Microcarbo 
ANWC BS-1419. Right: Phalacrocorax sulcirostris 
ANWCBS-1310. 

Holotype 

WAM 70.2.10, a pelvis with associated femora 
and caudal vertebrae. 



Discussion 

Australasia has four sympatric species of 
cormorants that occur on inland waters, which is 





1 








FIGURE 4. Caudal view of left femur. Microcarbo 
serventyorum WAM 70.2.10. 



more than anywhere in the world. Two are large, 
the Great cormorant, Phalacrocorax carbo, and 
the Pied cormorant, P. varius, and forage in rivers 
and large lakes and lagoons. Two are small, the 
Little black cormorant, P. sulcirostris, and the 
Little pied cormorant, Microcarbo melanoleucos . 
Phalacrocorax sulcirostris forages mainly in 
flocks in open water and M. melanoleucos forages 
in ponds, streams and along edges of large water 
bodies. Species of Phalacrocorax swim rapidly 
with their feet behind the body, while species of 
Microcarbo swim slowly with their feet beside 



TABLE 1 


. Measurements (mm) of pelves and femora of 


species of Microcarbo and Phalacrocorax. 


A: 


x (sd, 


range, n) 














M. melanoleucos 


P. sulcirostris 


P. olivaceus 



TL 


82(4,73-88, 13) 


97(6,84-113,25) 


102(7,95-109,3) 


WA 


25(1.4,23-28, 15) 


27(1.4,24-30,25) 


30(2.3,27-31,3) 


LF 


2.8(0.4,2.0-3.3, 14) 


3.2(0.3,2.6-3.9,25) 


3.3(0.4,2.9-3.6,3) 


PW 


11 (0.7,9-12, 14) 


12(0.8, 11-14,25) 


13(1.2, 12-14,3) 


DH 


5.0(0.3,4.5-5.5, 14) 


4.9 (0.3, 4.3-5.3, 25) 


5.6(0.6,4.9-6.0,3) 


TLAVA 


3.2(0.13,3.0-3.5, 13) 


3.6(0.18,3.2-38,25) 


3.4(0.17,3.2-35,3) 



B:n= 1 



M. qfricanus M. coronatus M. pygmaeus M. niger M. serventyorum 



TL 


79 


77 


81 


77 


71 


WA 


25 


26 


26 


23 


22 


LF 


2.5 


2.7 


2.7 


2.9 


5.1 


PW 


11 


11 


11 


9 


7.5 


DH 


4.7 


4.6 


5.0 


4.1 


4.3 


TLAVA 


3.2 


3.0 


3.1 


3.3 


3.2 



Abbreviations: TL = length of medium dorsal ridge of pelvis, WA = width between tips of antitrochanters, LF 
length of 8th postacetabular foramen, PW = proximal width of femur, and DH = depth of head of femur. 



138 



G. F. VAN TETS 



the body. Exaggeration of pelvic features that 
distinguish extant species of Microcarbo from 
those Phalacrocorax suggest that Microcarbo 
serventyorum was even more adept at foraging in 
confined bodies of water. Its small size and 
discovery in a peat swamp suggest that M. 
serventyorum lived in marshes with dense 
vegetation, small pools and narrow creeks. It may 
have had a niche distinct from that of the other 
four species of cormorant and may have been part 



of the Australasian divergence I postulated (van 
Tets 1976). 



Acknowledgments 

I thank Walter Boles, Nick Klomp and Trevor Worthy 
for their comments on the text, the Western Australian 
Museum for making the holotype available for study, 
Dragi Markovic for the photography and Carolyn Roach 
and Jo Heindl for typing the manuscript. 



References 



BONAPARTE, C. L. J. L. 1855 
Avium'. Vol. 2. Leiden. 



Conspectus Generum 



MARCHANT, S. & HIGGINS, P. J., eds. 1990. 
'Handbook of Australian, New Zealand and Antarctic 
Birds'. Vol. 1. Oxford University Press, Melbourne. 

OLSON, S. L. 1975. An evaluation of the supposed 
anhinga of Mauritius, Auk 92: 374-376. 

SERVENTY, D. L. 1938. The feeding habits of 
cormorants in south-western Australia. Emu 38: 293- 

316. 

SERVENTY, D. L. 1939. Notes on cormorants. Emu 38: 

357-371. 

SERVENTY, D. L., SERVENTY, V. & WARHAM, J. 
1971. The Handbook of Australian Sea-birds'. Reed, 

Sydney. 



SIEGEL-CAUSEY, D. 1988. Phylogeny of the 
Phalacrocoracidae. Condor 90: 885-905. 

VAN TETS, G. F. 1976. Australasia and the origin of 
shags and cormorants, Phalacrocoracidae. Pp. 121- 
124 in 'Proceedings of the 16th International 
Ornithological Congress'. Eds. H. J. Frith & J. H. 
Calaby. Australian Academy of Science, Canberra. 

VAN TETS, G. F. & O'CONNOR, S. 1983. The Hunter 
Island Penguin, an extinct new genus and species 
from a Tasmanian Midden. Records of the Queen 
Victoria Museum SI: 1-13. 

VAN TETS, G. F., RICH, P. V. & MARINO- 
HADIWARDOYO, H. R. 1989. A reappraisal of 
Protoplotus beauforti form the early Tertiary of 
Sumatra and the basis of a new pelecaniform family. 
Publication of the Geological Research and 
Development Centre, Paleontology Series 



BIRDS FROM THE BLUFF DOWNS LOCAL FAUNA, ALLINGHAM 

FORMATION, QUEENSLAND 

Walter E. Boles & Brian Mackness 



Summary 

A small number of bird fossils have previously been recorded from the early Pliocene freshwater, 
fluviate and lacustrine deposits of the Allingham Formation, nowrthwest of Charters Towers, 
northeastern Queensland. There has been significant avifaunal discoveries made during recent 
excavations of the site, most of these being watchbirds. The number of taxa now recognised has 
almost trebled from previous published accounts and includes at least seven orders, ten families and 
15 species-leval taxa. A new subspecies of the Purple swamphen, Porphyrio porphyrio nujagura 
subsp. nov., is described. The earliest occurrences of the genera Anhinga, Ardea, Cereopsis and 
Progura in the Australian avifaunal fossil record are also reported. The potential role that the study 
of fossil birds can play in palaeocological reconstruction is discussed. 



BIRDS FROM THE BLUFF DOWNS LOCAL FAUNA, ALLINGHAM FORMATION, 

QUEENSLAND. 



WALTER E. BOLES & BRIAN MACKNESS 



BOLES, W. E. & MACKNESS, B. 1994. Birds from the Bluff Downs Local Fauna, Allingham 
Formation, Queensland. Rec. S. Aust. Mus. 27(2): 139-149. 

A small number of bird fossils have previously been recorded from the early Pliocene 
freshwater, fluviatile and lacustrine deposits of the Allingham Formation, northwest of Charters 
Towers, northeastern Queensland. There has been several significant avifaunal discoveries made 
during recent excavations of the site, most of these being waterbirds. The number of taxa now 
recognised has almost trebled from previous published accounts and includes at least seven 
orders, ten families and 15 species-level taxa. A new subspecies of the Purple swamphen, 
Porphyria porphyria nujagura subsp. nov., is described. The earliest occurrences of the genera 
Anhinga, Ardea, Cereopsis and Progura in the Australian avifaunal fossil record are also 
reported. The potential role that the study of fossil birds can play in palaeoecological 
reconstructions is discussed. 

Walter E. Boles, Division of Vertebrate Zoology, Australian Museum, 6-8 College Street, 
Sydney. Brian Mackness, School of Biological Sciences, University of New South Wales, P.O. 
Box 1, Kensington New South Wales 2033. Manuscript received 31 July 1993. 



Birds have long been considered to be of 
secondary importance in the examination of fossil 
assemblages, with the paucity of their occurrence 
in the fossil record being cited as just one of the 
reasons for this cursory treatment (Olson 1985). 
The Australian fossil avifaunal record stretches 
back to the Cretaceous (Vickers-Rich 1991) and, 
although there has been a considerable increase in 
the investigation of fossil birds in recent years, 
most Tertiary studies have been taxonomic in 
nature, usually focussing on a single group (e.g. 
Miller 1963, 1966; Rich 1979; van Tets 1974). 
The interpretation of palaeoenvironments using 
fauna has relied almost exclusively on mammals, 
and indeed the 'Local Fauna' concept (Tedford, 
1970) has this as its foundation. 

In recent years, however, with an increasing 
number of palaeoornithologists working in 
Australia, birds are being used more and more 
both as biostrati graphic markers (Rich 1979) and 
as palaeoecological indictors (Boles 1993; Baird 
1993). 

A variety of taxa has been recovered from the 
early Pliocene freshwater fluviatile and lacustrine 
deposits of the Allingham Formation, northwest 
of Charters Towers, northeastern Queensland 
(Archer 1976, Bartholomai 1978, Archer & 
Dawson 1982, Archer 1982, Rich & van Tets 
1982). Collectively this assemblage has been 
called the Bluff Downs Local Fauna (Archer 
1976). The only bird mentioned in the original 



description of the fauna was Xenorhynchus 
[=Ephippiorhynchus] asiaticus (Archer 1976). 
Subsequently Rich & van Tets (1982) cited five 
taxa: Xenorhynchus cf. X. asiaticus, Threskiornis 
sp., cf. Dendrocygna sp., Numenius sp. and 
Charadriiformes. Rich et al. (1991) also listed five 
taxa: Xenorhynchus [=Ephippiorhynchus] 
asiaticus, Threskiornis sp., Cygnus sp., 
Dendrocygna sp. and Numenius sp. None of these 
listings offered further elaboration. 



Materials and methods 



Measurements were taken with vernier calipers 
accurate to 0.05 mm and rounded to the nearest 
0.1 mm. Terminology of bones is primarily from 
Baumel (1979). Where comparisons were made 
with published measurements, the methods of 
measuring followed those adopted in the 
comparative study; otherwise these largely 
followed those illustrated by Steadman (1980). 
Specimens are currently held at the Queensland 
Museum. All the fossil material examined and all 
modern specimens used for comparisons were 
considered to be adults, based on the absence of a 
'. . . pitted surface of the bone and incomplete 
ossification of the articular facets' (Campbell 
1979: 17). 



140 



W. E. BOLES & B. MACKNESS 




BLUFF DOWNS LOCAL FAUNA 



141 



Systematics 



ANHINGIDAE 



PHALACROCORACIDAE 

Phalacrocorax sulcirostris 

(Fig. la,b) 

Material 

Proximal right humerus (QM F23242), distal 
right humerus (QM F23241), possibly from the 
same bone. Measurements: proximal width 16.8 
mm, distal width 11.7 mm, depth of condylus 
dorsalis 7.9 mm. Locality: EVS Site. 

Characters 

The two fragments are considered to belong to 
the same species, and possibly the same 
individual, on the basis of size and configuration. 
The proximal end is referred to the 
Phalacrocoracidae on the basis of the reduced 
crista pectoralis, very broad impressio m. 
coracorbrachialis and broad, deep sulcus 
ligamentosus transversus. The distal fragment has 
a deeply excised fossa m. brachialis, which 
among extant Australian species occurs in P. 
sulcirostris and P. varius (Siegel-Causey 1988). 
The fossil agrees in size with P. sulcirostris 
(proximal width 16.2-17.2 mm, distal width 11.4- 
13.1 mm, depth of condylus dorsalis 7.9-8.6 
mm), and is smaller than P. varius (proximal 
width 19.1-21.8 mm, distal width 13.1-14.1 mm, 
depth of condylus dorsalis 9.1 - 9.8 mm). 

Remarks 

The Little Black Cormorant is widespread in 
wetlands, prefering open water greater than one 
metre deep, including large lakes, areas with 
flooded or fringing trees, and swamps with 
permanent or semi-permanent water (Marchant & 
Higgins 1990). 



Anhinga sp. 

Material 

Proximal right humerus (QM F23653); right 
carpometacarpus (QM F25776). This material 
represents a new species and is being described 
(B. M.) elsewhere. Locality: QM F23653, EVS 
Site; QM F25776, Main Site. 

Remarks 

Darters prefer smooth, open water at least 0.5 m 
deep, including permanent waterbodies, large 
lakes with shallow vegetated edges and semi- 
permanent swamps (Marchant & Higgins 1990). 



ARDEIDAE 

cf. Ardea picata 

(Fig. le) 

Material 

Distal right tibiotarsus (QM F23243). 
Measurements: distal width 7.7 mm, depth of 
condylus lateralis 1 A mm, depth of condylus 
medialis 7.9 mm. Locality: EVS Site. 

Characters 

The fossil is smaller and relatively more gracile 
than Egretta novaehollandiae, E: garzetta, A. 
intermedia, Ixobrychus flavicollis and Nycticorax 
caledonicus, but a good match in size and 
robustness for Ardea picata (distal width 6.3-8.0 
mm, depth of condylus lateralis 6.5-7.6 mm, 
depth of condylus medialis 6.6-7.9 mm). It is not 
possible to assign the distal tibiotarsus to any 
particular genus of herons on morphological 
features. Payne & Risley (1976) found no 
consistent differences in the tibiotarsi within this 
family. 



FIGURE 1. Avian fossils from the Bluff Downs Local Fauna. Bars equal 10 mm. Bars are shared by A and B, and by 
M,N and O. Queensland Museum registration numbers are given in parentheses. 

A. Phalacrocorax sulcirostris, proximal right humerus (F 23242); B. Phalacrocorax sulcirostris, distal right 
humerus (F 23241); C. Threskiornis sp. cf. T. molucca, cranial right coracoid (F 23257); D. Threskiornis sp. cf. T. 
molucca, distal right tarsometatarsus (F 23256); E. Ardea sp. cf. A. picata, distal right tibiotarsus (F 23243); F. 
Ephippiorhynchus asiaticus, proximal left humerus (F 23244); G. Ephippiorhynchus asiaticus, distal left humerus 
(F 23245); H. Phoenicopterus sp. cf. P. ruber, distal right tarsometatarsus (F 23252); I. Dendrocygna arcuata, 
proximal right humerus (F 23248); J. Progura sp. cf. P. naracoortensis, carpometacarpus (F 23258); K. Progura 
sp. cf. P. naracoortensis, proximal left tarsometatarsus (F 23259); L. Porphyria porphyrio nujagura subsp. nov., 
proximal right tarsometatarsus (F 23250); M. Rallidae gen. and sp. indet. 2, distal left tarsometatarsus (F 23255); N. 
Rallidae gen. and sp. indet. 1, distal left tibiotarsus (F 23254); O. Porzana sp., distal left tarsometatarsus (F 23253); 
P. cf. Numenius sp., distal right femur (F 23251). 



142 



W. E. BOLES & B. MACKNESS 



Remarks 

Virtually all herons are associated with water, 
the type of wetland preferred and the manner in 
which it is utilised depending on the species. The 
Pied Heron occurs in shallow wetlands, including 
floodplains and swamps (Marchant & Higgins 
1990). 



CICONIIDAE 

Ephippiorhynchus asiaticus 

(Fig. lf,g) ' 

Material 

Proximal left humerus (QM F23244); distal left 
humerus (QM F23245). Measurements: proximal 
width 44.8 mm, distal width 34.4 mm, depth of 
condylus dorsalis 18.7 mm. A '. . . fragment of 
tarsometatarsus (QM F7036)' cited by Archer 
(1976: 385) was not examined. Locality: QM 
F23244 AB Site; QM F7036, QM F23245 Main 
Site. 

Characters 

The proximal fragment agrees with the 
Ciconiidae and differs from the Gruidae, the only 
other similar taxon, by having caput humeri 
proportionally shorter, intumescentia less inflated 
and fossa pneumotricipitalis larger. The distal 
fragment resembles the former family, but not the 
latter, by having tuberculum supracondylare 
ventrale narrow and ridge-like, and processus 
supracondylar is dorsalis prominently produced. 

Remarks 

The Black-Necked Stork, Australia's only 
extant member of this family, inhabits mainly 
open water up to 0.5 m deep, including extensive 
sheets over grassland, shallow swamps and pools 
on floodplains (Marchant & Higgins 1990). 



c. 8.0 mm, dorsal length of trochlea metatarsi HI 
8.2 mm. Locality: QM F23256 EVS Site; QM 
F23257 Main Site. 

Characters 

The tarsometatarsus is assigned to Threskiornis 
rather than Platalea because trochlea metatarsi II 
agrees with the former in being less recessed 
plantarly relative to trochlea metatarsi III (in 
medial view). It agrees in size with molucca 
(depth of trochlea metatarsi III 8.4 mm, plantar 
length of trochlea metatarsi HI 8.0 mm), which is 
larger than spinicollis (depth of trochlea metatarsi 
III 1 .3-7 .4 mm, plantar length of trochlea 
metatarsi HI 7.1-7.3 mm). 

Allocation of the coracoid to Threskiornis rather 
than Platalea is on the basis of having a narrower 
processus acrocoracoideus (in dorsal view), 
cotyla scapularis proportionally further from 
processus acrocoracoideus , and processus 
procoracoideus curving less mediad. 

Living T. spinicollis and T. molucca are not 
separable on measurements that can be taken on 
the fossil coracoid (cranial end of processus 
acrocoracoideus to cotyla scapularis: spinicollis 
20.4-21.8 mm, molucca 20.2-22.8 mm; depth of 
processus acrocoracoideus: spinicollis 11.7-12.1 
mm, molucca 1 1.3-12.1 mm). These species differ 
somewhat in robustness of the shaft; however, 
damage to the Bluff Downs material precludes 
comparison. The coracoid is referred to the same 
taxon as the tarsometatarsus. Because material is 
limited and the living species are morphologically 
quite similar, a more definite identification is not 
made. 

Remarks 

The Australian White Ibis habitat preferences 
include shallow water over soft substrates, in 
swamps and open water, and muddy flats 
(Marchant & Higgins 1990). 



THRESKIORNITHIDAE 

Threskiornis sp. cf. T. molucca 

(Fig. 1 c,d) 

Material 

Cranial right coracoid (QM F23257); distal 
right tarsometatarsus missing trochlea metatarsi 
IV (QM F23256). Measurements: coracoid, 
cranial end of processus acrocoracoideus to 
caudal end of cotyla scapularis 20.6 mm, depth of 
processus acrocoracoideus 9.4 mm; 
tarsometatarsus, depth of trochlea metatarsi HI 



PHOENICOPTERIDAE 

Phoenicopterus sp. cf. P. ruber 

(Fig. lh) 

Material 

Partial distal right tarsometatarsus lacking 
trochlea metatarsi IV (QM F23252). 
Measurements: medial depth of trochlea 
metatarsi II c. 7.9 mm, lateral depth of trochlea 
metatarsi II 8.4 mm, dorsal width of trochlea 
metatarsi HI 10.3 mm, lateral depth of trochlea 



BLUFF DOWNS LOCAL FAUNA 



143 



metatarsi HI c. 9.1 mm, dorsal width of trochlea 
metatarsi HI c. 7.6 mm, plantar length of trochlea 
metatarsi HI 1 1 .7 mm. Locality: Main Site. 

Characters 

This specimen is identified as a flamingo on the 
basis of the following characters (most from Rich 
et al. 1987): trochlea metatarsi II short relative to 
trochlea metatarsi HI; distal end of trochlea 
metatarsi II broader than plantar border; trochlea 
metatarsi II twisted plantarly and laterally from 
front of tarsometatarsus; and trochlea metatarsi 
III narrow and deep. Reference of this fragment to 
this species is made on the basis of size rather 
than on diagnostic morphology. 

The measurements of the Bluff Downs 
specimen are a good match for those of extant P. 
ruber given by Rich et al. (1987), and the 
specimen is therefore tentatively assigned to this 
taxon. This follows the practice of Rich et al. 
(1987: 207), who, faced with limited and 
undiagnostic material, segregated the forms on 
size and "... provisionally retained the generic 
and specific names that have priority as a 
convenience until more complete material allows 
a better evaluation of the systematic positions of 
the Pliocene and Quaternary flamingoes of 
Australia'. 

Remarks 

There are few records of flamingoes in Australia 
away from the centre of the continent. Rich et al. 
1991 have recorded an undetermined flamingo 
from the late Miocene Alcoota Local Fauna, 
northern Australia. Living P. ruber of Africa 
frequents mainly saline or alkaline lakes, estuaries 
and lagoons, seldom alighting on fresh water 
(Brown et al. 1982). Similar conditions have been 
proposed for central Australian lake deposits 
yielding flamingo remains. The Alcoota Locality 
during the late Miocene is considered by Murray 
& Megirian (1992: 214) to have represented '. . . a 
small but permanent, possibly spring-fed pond or 
lake, sometimes expanding to a temporary, large, 
shallow lake.' 



ANATIDAE 

Cygnus atratus 

Remarks 

This taxon was cited by Vickers-Rich (1991) 
without further elaboration. This material has not 
been examined in this study. Locality: Main Site. 



Cereopsis sp. 

Material 

A proximal carpometacarpus fragment (QM 
F23260). This material, probably representing the 
extant species C. novaehollandiae, will be 
described (B.M.) elsewhere. Locality: EVS Site. 

Remarks 

This species inhabits grasslands and terrestrial 
wetlands, occasionally entering water. 



Dendrocygna arcuata 

(Fig. li) 

Material 

Proximal right humerus (QM F23248). 
Measurements: proximal width c. 18.3 mm, depth 
of caput humeri 6.0 mm. Locality: Main Site. 

Characters 

The Anserinae and Dendrocygninae are 
separated from other Anatidae by the combination 
of prominent capital shaft ridge directed towards 
the caput humeri, attachment for caput ventrale 
of M. humerotriceps extending virtually to caput 
humeri, and area of attachment of M. pectoralis 
on the tuberculum dorsale is elevated and 
somewhat circular (Woolfenden 1961). The 
Anserinae is unrepresented in Australasia except 
by the aberrant Cereopsis, which is diagnosable 
by several unique characters. The Bluff Downs 
specimen is an excellent fit for Dendrocygna, 
agreeing in size with D. arcuata, the smaller of 
the two Australian species. 

Remarks 

The Water Whistling-Duck prefers fresh, deep 
permanent waters with emergent vegetation 
(Marchant & Higgins 1990). 



Nettapus sp. 

Material 

An almost complete left carpometacarpus (QM 
F23249). This material probably represents a new 
species and will be described (B.M.) elsewhere. 
Locality: EVS Site. 

Remarks 

Pygmy-geese are wholly aquatic on terrestrial 
wetlands, preferably deep (greater than lm), 
permanent water bodies, with abundant floating 



144 



W. E. BOLES & B. MACKNESS 



and submerged vegetation (Marchant & Higgins 
1990). 



MEGAPODIIDAE 

Progura sp. cf. P. naracoortensis 

(Fig. lj,k) 

Material 

Carpometacarpus (QM F23258) lacking os 
metacarpale minor. Proximal left tarsometatarsus 
(QM F23259) lacking most of the hypotarsus. 
Measurements: carpometacarpus, proximal width 
(dorsal) 17.5 mm, (ventral) 15.0 mm; 
tarsometatarsus, proximal width 19.5 mm. 
Locality: EVS Site. 

Characters 

A tentative identification has been made on size. 
Van Tets (1974) published measurements for the 
two known species: proximal width of 
tarsometatarsus, P. naracoortensis 21-23 mm, P. 
gallinacea 26-29 mm; dorsal and ventral widths 
of carpometacarpus, P. gallinacea 27 x 16 mm; a 
carpometacarpus of P. naracoortensis was not 
available. The Bluff Downs material is smaller 
and older than that of either species. Here it is 
tentatively referred to the smaller P. 
naracoortensis, but it may represent an 
undescribed species. 

The tarsometatarsus of Progura is separated 
from that of extant genera of Australian 
megapodes by the following combination of 
characters: hypotarsus situated more laterad, 
sulcus hypotarsi more laterally situated relative to 
eminentia inter condylaris ', cotyla medialis nearly 
size of cotyla lateralis in cranial view with dorsal 
border rounded and projecting dorsally about the 
same extent as cotyla lateralis, medial border of 
corpus tarsometatarsi straight (less concave), and 
sulcus extensorius broader and deeper and 
extending further distad. 

The carpometacarpus is distinguished by its 
combination of processus extensorius more 
proximally directed, proximal border of fades 
articularis radiocarpalis of trochlea carpalis 
rounded (not pointed), caudal border of fades 
articularis ulnocarpalis of trochlea carpalis 
rounded (not flattened) and extended ventrally 
only slightly more than fades articularis 
radiocarpalis, and os metacarpale majus slightly 
caudally curved. 

Remarks 

Previous records of Progura are from coastal 



and subcoastal areas of eastern and southeastern 
Australia. Not enough is known about the ecology 
of these animals for them to be useful 
bioecological indicators. Van Tets (1974) 
hypothesised that, based on relative leg lengths, 
the longer-legged P. gallinacea was a rainforest 
species, whereas P. naracoortensis inhabited open 
scrub land. Van Tets (1984) later suggested that, 
because these two species are usually found 
together, they could represent a single, sexually 
dimorphic species. 



RALLIDAE 

Porzana sp. 

(Fig. lo) 

Material 

Distal left tarsometatarsus (QM F23253). 
Measurements: distal width 4.3 mm. Locality: 
EVS Site. 

Characters 

This specimen is larger than Porzana tabuensis 
(distal width 3.7 mm) or P. pusilla (3.3 mm) and 
about the same size as P. fluminea (4.3 mm) and 
P. cinereus (4.7 mm). Compared with these latter 
two species, trochlea metatarsi are thinner and 
more splayed, trochlea metatarsi II appears 
slightly less directed plantarad, and incisurae 
intertrochlearis are wider. The specimen is 
abraded on the trochlear surfaces, and some of 
these differences could be artefacts of this wear. 
Osteological characters given by Olson (1970) and 
Steadman (1986) between P. cinerea and other 
species of Porzana are not relevant to this partial 
element. 

Remarks 

Most rails are associated with water. The 
smaller species, including most Porzana, are 
usually shy, spending most of their time in dense 
waterside vegetation. Porzana cinerea also 
requires floating vegetation. 



Genus and species indet. 1 

(Fig. In) 

Material 

Distal left tibiotarsus (QM F23254). Distal 
width 5.1 mm, depth of condylus lateralis c. 5.3 
mm, depth of condylus medialis c. 5.3 mm. 
Locality: EVS Site. 



BLUFF DOWNS LOCAL FAUNA 



145 



Remarks 

The fossil represents a small to medium rail 
between the sizes of Porzana cinereus (distal 
width 4.2 mm, depth of condylus lateralis 3.9 
mm, depth of condylus medialis 4.1 mm) and 
Gallirallus philippensis (distal width 6.0 mm, 
depth of condylus lateralis 6.0 mm, depth of 
condylus medialis 6.0 mm). There are several 
extant Australasian genera within this size range 
with which the specimen should be compared, 
Dryolimnas (sensu Olson 1973), Rallina and 
Rallicula. 



Genus and species indet. 2. 

(Fig. lm) 

Material 

Distal left tarsometatarsus (QM F23255). 
Measurements: distal width 7.1 mm, depth of 
trochlea metatarsi HI 3.8 mm. Locality: EVS Site. 

Characters 

This specimen comes from a larger and 
somewhat more robust species than the previous 
indeterminate rail. It is similar in size to 
Amaurornis olivacea (distal width 6.7-6.9 mm, 
depth of trochlea metatarsi HI 3.2-3.6 mm) and 
Gallinula ventralis (distal width 7.4 mm, depth of 
trochlea metatarsi HI A3 mm). This fossil is also 
similar in morphology to A. olivacea, and 
probably could be tentatively referred to that 
species; however, it seems prudent to await more 
extensive material and comparisons with a greater 
range of genera before taking this action. 



Porphyrio porphyrio nujagura subsp. nov. 
(Fig. 11) 



There are four Recent species of Porphyrio from 
Australasia: three flightless Australasian species 
(mantelli, New Zealand; albus, Lord Howe Island 
- extinct; and kukwiedei, New Caledonia - 
extinct; Balouet & Olson 1989); and porphyrio, 
the only member of the genus now occurring in 
Australia. The first three are much larger and 
robust than porphyrio. The Bluff Downs specimen 
shows no morphological differences from 
porphyrio but is smaller than either sex of this 
sexually size dimorphic species (Australian 
porphyrio: proximal width 10.7-12.5 mm, 
proximal depth 12.4-13.7 mm, length of 
hypotarsus 11.0-13.0 mm, width of hypotarsus 
6.2-6.9 mm). Compared with measurements given 
by Steadman (1988: Table 2), it is also smaller 
than most extralimital populations of the P. 
porphyrio superspecies, except P. porphyrio from 
Bechuanaland and P. poliocephalus of Thailand. 

Etymology 

The specific name is from the Gugu-Yalanji 
dialect word nujagura, meaning 'prehistoric 
times' (Oatesefa/. 1964). 

Remarks 

The only extant Australian species of 
comparable size not examined was Eulabeornis 
castaneoventris, a mangrove specialist, for which 
no skeletons exist. The end of the shaft is jagged, 
indicating that the break occurred before 
fossilisation.This individual was probably a victim 
of a predator or scavenger. The Purple swamphen 
prefers permanent freshwaters with good cover of 
rushes and other larger waterplants, at least along 
the water's edge, usually in the proximity of more 
open grazing areas (Marchant & Higgins 1993). 



SCOLOPACIDAE 



Material 

Proximal right tarsometatarsus (QM F23250). 
Measurements: proximal width 9.7 mm, proximal 
depth 11.4 mm, length of hypotarsus 10.4 mm, 
width of hypotarsus 5.6 mm. Locality: EVS Site. 

Characters 

Agrees with Porphyrio and differs from other 
genera of Australian rails, including the genera of 
larger forms (Gallinula, including Tribonyx; 
Fulica; Gallirallus australis) by having a distally 
directed projection on the plantodistal end of the 
hypotarsus (in other genera, the hypotarsus curves 
smoothly into the shaft). 



cf. Numenius sp. 

(Fig. lp) 

Material 

Distal right femur (QM F23251). 
Measurements: distal width 10.0 mm, depth of 
condylus lateralis 8.7 mm, depth of condylus 
medialis c. 7.0. Locality: Main Site. 

Characters 

This specimen appears to represent a large 
sandpiper. It is substantially larger than all 
Australasian taxa except for Numenius, for which 
it is also a good match in morphology. Because 



146 



W. E. BOLES & B. MACKNESS 



the distal femur has limited diagnostic value and 
the bone is slightly abraded, identification is not 
attempted beyond cf. Numenius sp. 

Remarks 

Some members of this family are restricted to 
coastal areas; others occur in freshwater wetlands 
(Lane 1987). Larger Numenius species are coastal, 
whereas N. minutus extends well into subcoastal 
regions. 



Discussion 



Taphonomy 



All fossils were recovered from a series of 
massive lacustrine clays from Main Quarry 
(Archer 1976) and Elaine's Vertebrae Site 
(Mackness unpublished information). Most of the 
bones were not complete and showed post- 
depositional breakage and fragmentation. An 
exception was the tarsometatarsus of Porphyria 
porphyria nujagura, the jagged edge of which 
suggested predation or scavenging while the body 
was still relatively fresh. There was little evidence 
of transportation wear and, even though there was 
no articulation, it is probable that the birds died in 
reasonable proximity to the site of deposition. An 
examination of bone textures suggests that there 
was little subaerial prediagenetic exposure and 
that most bones were quickly buried or 
submerged. 

Vickers-Rich (1991) suggested a bias in 
avifaunal fossil deposits toward medium-sized 
birds (e.g. ducks, flamingos, burhinids) and larger 
birds (emus) to the exclusion of smaller birds. 
Although the bird assemblage recovered to date is 
consistent with this prediction, large scale wet- 
screening of sediments presently being undertaken 
may result in the recovery of smaller birds. The 
depositional environment obviously favours the 
preservation of waterbirds: these are also common 



Coracoid 


■ 


(i) 




Humerus 






(4) 










Carpometacarpus 






(4) 










Femur 




■ (2) 












Tibiotarsus 




■ (2) 












Tarsometatarsus 









(5) 



FIGURE 2. Summary of avian skeletal elements 
recovered from Bluff Downs Site. 



in central Australian Tertiary sites (Vickers-Rich 
1991). The proportions of the different bone 
elements found fossilised at Bluff Downs are 
summarised in Figure 2. All are regarded to be the 
most durable elements (Napawongse 1981; Rich 
&Baird 1986; Vickers-Rich 1991). 



Palaeoecology 

Archer (1976) has suggested that the Bluff 
Downs Local Fauna may have been riparian. The 
large number of non-avian fossils recovered from 
the Bluff Downs site include both terrestrial and 
aquatic forms, with neither predominating. The 
mammals provide no definitive indication of what 
the terrestrial environment may have been like, 
although a typically rainforest-dwelling 
pseudocheirid possum is presently being described 
(B.M), along with an enigmatic family of 
marsupials (Mackness et al. 1993). 

Three types of crocodiles have been recovered, 
including two large aquatic and one terrestrial 
form (Willis & Mackness in prep.). Studies of the 
molluscan fauna have revealed a diverse suite of 
species, some requiring specific aquatic niches 
that range from high energy fluvatile environments 
to stagnant lacustrine regimes (Mackness, 
unpublished data). It is evident from this faunistic 
'mix' that there was a complex series of aquatic 
environments available either ephemerally or on a 
permanent basis. 

Various authors (Frith 1959, Braithwaite & 
Frith 1969, Braithwaite 1975, Fjeldsa 1985) have 
suggested a relationship between the distribution 
of waterbird species and habitat types. Fjeldsa 
(1985) found poor correlation between avian 
communities and classifications based on 
vegetation types. Indicator species, e.g. Porphyria 
and Phalacrocorax, are present in several of 
Fjeldsa's classifications. Given the taphonomic 
evidence of minimal transportation and the nature 
of the sediments themselves, the Bluff Downs 
avifauna appears to be a biocoenotic assemblage, 
which may represent several depositional 
episodes. While most of the taxa identified, or 
their closest extant relatives, are relatively 
nonspecific in their preferences for wetland types, 
some utilise a range of microhabitats within 
wetland environments. Living species of Nettapus 
are more strict in their requirements than most, 
being generally confined to deeper water with 
considerable floating vegetation. 

Living Cereopsis novaehollandiae graze on 
grasslands, whereas living phoenicopterids are 



BLUFF DOWNS LOCAL FAUNA 



147 



TABLE 1 . Comparison of Australian Pliocene avifaunal assemblages. Data for Chinchilla and Kanunka from Rich el 

al. ( 1 99 1 ) and Vickers-Rich ( 1 99 1 ). 



Family 



Bluff Downs 



Casuariidae 


- 


Pelecanidae 


- 


Phalacrocoracidae 


X 


Anhingidae 


X 


Ardeidae 


X 


Ciconiidae 


X 


Threskiornithidae 


X 


Phoenicopteridae 


X 


Anatidae 


X 


Accipitridae 


- 


Gruidae 


- 


Megapodidae 


X 


Rallidae 


X 


Otididae 


- 


Scolopacidae 


X 


'Charadriiformes' 


- 



Chinchilla 



X 



Kanunka 



X 
X 
X 
X 
X 
X 

X 
X 
X 
X 



normally found in saline environments today. 
Disregarding these novelties, the remaining taxa 
comprise a waterbird community that differs little 
from that which now occurs in tropical Australia, 
such as in the wetlands of Kakadu National Park, 
Northern Territory, an area supporting seasonal 
floodplains, waterholes, rivers, ephemeral swamps 
and permanent lakes. From the avian assemblage, 
it is probable that at least part of Bluff Downs 
consisted of similar wetlands during the Pliocene. 



Comparisons with other Pliocene faunas 

Most Pliocene deposits have yielded bird 
fossils. Many are too fragmentary for identification 
while others have not yet been studied (Vickers- 
Rich 1991). Apart from the Bluff Downs Local 
Fauna, both the Chinchilla and Kanunka Local 
Faunas have significant avian components (Table 
1). All three avifaunas are considered to be 
wetland assemblages, with the dominant families 
represented containing mostly wetland specialists. 
In these, most of the same families are 
represented. However long-legged wading birds 



(Ciconiiformes, Phoenicopteriformes) are absent 
from Chinchilla. 



Acknowledgments 

The authors wish to thank Michael Archer and 
Suzanne Hand (University of New South Wales) for 
providing helpful comments on the manuscript. Les 
Christidis and Rory O'Brien (Museum of Victoria), Jerry 
van Tets, (CSIRO Wildlife and Ecology), and Ralph 
Molnar (Queensland Museum) made specimens and/or 
comparative material available. Figure 1 was 
photographed and produced by Maurice Ortega, Stuart 
Humphreys, Lynne Albertson and Sally Cowan. Jack, 
Rhonda, Bram, Troy and Selesti Smith of Bluff Downs 
Station provided vital help and support for the ongoing 
research into the Bluff Downs Local Fauna. The 
collection of the Bluff Downs material was supported in 
part by: an ARC Program Grant to M. Archer; a grant 
from the Department of Arts, Sport, the Environment, 
Tourism and Territories to M. Archer, S. Hand and H. 
Godthelp; a grant from the National Estate Program 
Grants Scheme, Queensland to M. Archer and A. 
Bartholomai; and grants in aid to the Riversleigh 
Research Project from Wang Australia, ICI Australia and 
the Australian Geographic Society. 



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MIMICRY IN ANKYLOSAURID DINOSAURS 

Tony Tholborn 

Summary 

The tail club of Late Cretaceous ankylosaurid dinosaurs is usually regarded as a weapon to deter 
predators. However, the effective reach of the club was clearly constrained by the shortness and 
limited flexibility of the tail as a whole. Analogies with the defensive adaptations of butterflies and 
other insects indicate that the ankylosaurid tail club may represent a 'dummy head' that diverted 
predators from the true head. Simulated escape movements of that false head could have elicited the 
attack reponse of persistent aggressors, thus bringing them within striking range; thereupon the tail 
club would have performed its third function, as a weapon, to greatest effect. In performing such 
deceptive functions, the ankylosaurid tail club would qualify as an example of mimicry, seemingly 
the first to be identified among dinosaurs. 



MIMICRY IN ANKYLOSAURID DINOSAURS 



TONY THULBORN 



THULBORN, T. 1994. Mimicry in ankylosaurid dinosaurs. Rec. S. Aust. Mus. 27(2): 151-158. 

The tail club of Late Cretaceous ankylosaurid dinosaurs is usually regarded as a weapon to 
deter predators. However, the effective reach of the club was clearly constrained by the shortness 
and limited flexibility of the tail as a whole. Analogies with the defensive adaptations of 
butterflies and other insects indicate that the ankylosaurid tail club may represent a 'dummy 
head' that diverted predators from the true head. Simulated escape movements of that false 
head could have elicited the attack response of persistent aggressors, thus bringing them within 
striking range; thereupon the tail club would have performed its third function, as a weapon, to 
greatest effect. In performing such deceptive functions, the ankylosaurid tail club would qualify 
as an example of mimicry, seemingly the first to be identified among dinosaurs. 

Tony Thulborn, Vertebrate Palaeontology Laboratory, Department of Zoology, University of 
Queensland, Brisbane, Queensland 4072. Manuscript received 21 April 1993. 



Ornithischians of the suborder Ankylosauria are 
often epitomized as 'armoured dinosaurs' or 
'reptilian tanks' on account of their exuberant 
developments of dermal bone (e.g. Chang 1979; 
Norman 1985; Carroll 1988; Coombs & 
Maryanska 1990). Among these dinosaurs the 
skull was protected by a helmet-like covering of 
osteoderms, while the neck, back, flanks and tail 
were shielded by a remarkable array of bony studs 
and plates, often elaborated into keels or spikes 
(Fig. 1). In addition, the members of one 
ankylosaur group, the family Ankylosauridae, 
carried a conspicuous deterrent weapon in the 
form of a bony club at the tip of the tail (Fig. 2). It 
is often supposed that ankylosaurids would sweep 
this weapon sideways to strike at the feet and legs 
of their aggressors (e.g. Charig 1979; Coombs 
1979; Norman 1985). 

Conventional thinking on the defensive 
capabilities of the armoured dinosaurs has been 
well summarized by Coombs & Maryanska (1990: 
482-3): 

'When attacked by predators, ankylosaurs primarily 
defended themselves passively, relying on their 
extensive armor and low-slung, difficult-to-overturn 
body conformation. Outrunning of predators seems 
unlikely. Ankylosaurids may have used their tail 
clubs for active defense by sweeping it just above the 
ground to strike at the fragile ankles of an attacking 
predator...' 

This paper examines more closely the defensive 
capabilities of the ankylosaurid dinosaurs, giving 
particular attention to the role of the tail club. 



The ankylosaurid tail club 

The deterrent value of a tail club depends on its 
effective reach and, hence, on the length and 
flexibility of the entire tail. Surprisingly, the 
ankylosaurid tail was rather short and somewhat 
inflexible (Fig. 2). For instance, the ankylosaurid 
Euoplocephalus tutus possessed only 20 caudal 
vertebrae (excluding the first, which was joined to 
the sacrum, and an unknown number built into the 
terminal club); by comparison, the clubless tail of 
the nodosaurid Sauropelta edwardsi (family 
Nodosauridae) comprised 40 to 50 vertebrae 
(Carpenter 1982, 1984). Long transverse processes 
probably imposed some constraint on flexures at 
the base of the tail, and the distal caudal vertebrae 
were deeply internested, sheathed in ossified 
tendons and amalgamated into a rigid 'handle' to 
the tail club (Coombs & Maryanska 1990; 
Carpenter 1982). Consequently, the tail was 
flexible only in its proximal half, and then to a 
limited degree. It is inconceivable that the tail club 
could have been swung alongside the animal's 
flank, or across its back, or anywhere near the 
head, neck and shoulders. 

It is sometimes implied that special defensive 
behaviour might have compensated for the limited 
reach of the tail club. For example, Norman 
supposed (1985: 168) that the ankylosaurid 
Euoplocephalus was sufficiently agile to avoid the 
lunges of a tyrannosaur while manoeuvring into 
position to retaliate. However, the exceptionally 
broad body of ankylosaurs probably conferred 
great stability (Carpenter 1984; Norman 1985; 



152 



T. THULBORN 





FIGURE 1. General body form of ankylosaurian dinosaurs, a, Restoration of the nodosaurid ankylosaur Sauropelta 
edwardsi in dorsal view (adapted from Carpenter 1984). b, Comparative restoration of the ankylosaurid ankylosaur 
Euoplocephalus tutus (adapted from Carpenter 1982). Each scale bar indicates 1 metre. 



Bakker 1986; Coombs & Maryanska 1990), so 
that these animals were not easily overturned by 
predators. Such inherent stability is the antithesis 
of agility - which, in essence, is controlled 
instability. It is difficult to imagine that a single 
ankylosaurid, however agile, could turn rapidly 
enough to fend off two or more predators hunting 
in cooperation. 

Overall, it seems as if the ankylosaurid tail club 
could have functioned as an effective weapon only 
if an aggressor strayed within the rather limited 
reach of the tail. 



Behaviour of predatory dinosaurs 

Big theropod dinosaurs, or 'carnosaurs 1 sensu 
lato, were probably the only predators capable of 
killing and dismembering the heavily-armoured 
ankylosaurids, which grew to a length of 6 metres 
and a weight of several tonnes. Presumably, 
carnosaurs attacked the most vulnerable regions of 
their prey, namely the head and neck. This strategy 
has several advantages (Ewer 1968): an attack to 
the head may confuse and disorient the prey, 
thereby reducing the risk of retaliation; there is 



great likelihood of inflicting fatal injuries — by 
severing or puncturing the spinal cord, the trachea 
or major blood vessels; and, finally, a rapid kill 
reduces the risk of attracting competitors and 
scavengers. A tendency to bite the neck or head of 
the prey is widespread among existing predators, 
including a variety of mammalian carnivores 
(Ewer 1968, 1973; Eaton 1970; Schaller 1972; 
Leyhausen 1973) and birds (Cade 1967; 
Scherzinger 1970; Ullrich 1971; Smith 1973), and 
experiments with hand-reared animals indicate 
that this may be an innate response (Leyhausen 
1973; Lorenz & Leyhausen 1973; Smith 1973). 
The most important visual cues eliciting that 
response appear to be shape and motion of the 
prey. With respect to shape, experimental findings 
indicate that mammalian and avian predators tend 
to bite the most obvious constriction of the prey's 
body — which is usually the region of the neck. 
With respect to movement, those predators tend to 
bite the 'leading' end of the prey — which is 
usually the head. 

Palaeobiological evidence surveyed by Molnar 
& Farlow (1990) does not rule out the possibility 
that carnosaurs used prey-killing techniques 
comparable to those of existing big cats, which 



MIMICRY IN DINOSAURS 



53 




FIGURE 2. Tail skeleton of the ankylosaurid dinosaur Euoplocephalus tutus, a, Left lateral view, b, Dorsal view. 

Scale bar indicates 50 cm. 



frequently employ a bite to the head or neck 
(Schaller 1967, 1972; Ewer 1973). This possibility 
does not imply that all carnosaurs adhered to a 
single pattern of prey-killing behaviour; rather, a 
tendency to seize the head or neck may be 
envisaged as a basic technique, which could have 
been modified (or even abandoned) to suit the 
circumstances of diverse carnosaurs. 

It is noteworthy that many of the quadrupedal 
ornithischians susceptible to carnosaur attack 
(stegosaurs, ankylosaurids and nodosaurids) have 
a shallow skull that merges insensibly into a short 
neck. In effect, all these slow-moving herbivorous 
dinosaurs seem to have rendered the skull and 
neck as inconspicuous as possible. Among 
ankylosaurs there was also a tendency for the head 
and neck to be protected by a remarkable array of 
defensive structures. The head was encased in a 
veritable helmet of osteoderms, while the 
vulnerable region of the eye was shielded by a 
bony eyelid (Coombs 1972) and by an 
overhanging cornice of the skull roof. The shadow 
cast by this cornice may have concealed the eye, 
thus serving much the same role as a pigmented 
eye-stripe in many living animals (Cott 1940). In 
some ankylosaurs, such as the nodosaurid 
Edmontonia rugosidens, the neck was protected 
not only by its own covering of osteoderms but 
also by prominent bony spikes extending forwards 
from the region of the shoulder (Carpenter 1990, 
fig. 21.4; Coombs & Maryanska 1990, fig. 22.13). 
The existence of these elaborate defensive 
structures seems to confirm that the head and neck 
were particularly vulnerable regions of the 
ankylosaur body. 

Theoretically, it is possible to test the 
suggestion that carnosaurs tended to attack the 



head and neck of their prey: such behaviour might 
be expected to result in an unusually high 
incidence of teeth-marks on the neck and skull 
bones of animals killed by carnosaurs. 
Unfortunately, several factors conspire to render 
this test impracticable. First, it seems that 
predatory dinosaurs left surprisingly few teeth- 
marks on the bones of their prey (Fiorillo 1991). 
Second, teeth-marks in regions other than the head 
and neck would result from predators and 
scavengers dismembering carcasses and stripping 
them of their flesh. Next, some teeth-marks would 
be lost if bones were swallowed and partly or 
wholly digested by carnosaurs (Fiorillo 1991). 
And, finally, it is possible that the head of the prey 
was consumed preferentially, as among domestic 
cats and other small carnivores (Leyhausen 1973; 
Ewer 1968). For those reasons, the distribution 
and abundance of tooth-marks is unlikely to 
provide any clear indication to the prey-killing 
behaviour of carnosaurs. Moreover, it is 
unrealistic to assume that all carnosaurs shared a 
single pattern of prey-killing behaviour; instead, 
they probably exploited a repertoire of killing 
techniques (Molnar & Farlow 1990), each 
appropriate to the identity, size, weaponry and 
behaviour of particular prey animals. An 
analogous situation exists among mammalian 
carnivores, where an extensive range of prey- 
killing techniques appears to have been developed 
from the basis of a straightforward neck-bite 
(Ewer 1973). 



Heads and tails 
The heads and tails of ankylosaurids show 



154 



T.THULBORN 





FIGURE 3. Silhouettes of dinosaurs, to illustrate 
potential role of ankylosaurid tail club as a 'dummy 
head', a, The ankylosaurid dinosaur Euoplocephalus 
tutus (with head at right); feet and tip of snout are 
truncated, as if concealed by low vegetation, and the tail 
is raised, b, The ornithopod dinosaur Iguanodon in 
quadrupedal posture (with head at left); tail is truncated, 
as if lying on the ground or concealed by vegetation. 



divergent modifications in form: the head was 
rendered as inconspicuous as possible whereas the 
visual impact of the tail was exaggerated by the 
terminal club of bone. The net result is that head 
and neck resemble a smoothly tapering tail, 
whereas the tail has much the profile of a long 
neck and prominent head (Fig. 3). This unusual 
body outline calls to mind those insects that carry 
a conspicuous 'dummy head 1 at the rear end of the 
body (Fig. 4). Such dummy heads appear to divert 
predators away from the true head and towards 
the least vital extremity of the insect's body (Cott 
1940; Wickler 1968; Edmunds 1974; Curio 1976). 
It is suggested here that the ankylosaurid tail club 
was an analogous dummy head that served to 
deceive predatory dinosaurs. 

Before pursuing that suggestion, it must be 
noted that the term 'dummy head' (or 'false head') 
seems to have been applied indiscriminately to 
two rather different adaptations among living 
animals. Here I propose to separate 'dummy 
heads' into two categories — 'duplicate' heads and 
'substitute' heads. In some animals the head and 
tail are virtually identical in shape, as among 
amphisbaenians, snakes, typhlopids and certain 
scincid lizards. Here the tail duplicates the form of 
the head, or vice versa (Wickler 1968), thereby 
confusing and disorienting predators (Wickler 
1968; Bustard 1969). In ankylosaurids, and their 
insect analogues, the tail appears to be modified 
as a visual substitute for the head (and vice versa), 
evidently serving to draw the attention of predators 
away from the true head. Here the substitute head 



is more conspicuous than the true head. Although 
substitute heads may have originated through the 
elaboration of duplicate heads, the two adaptations 
are in many cases sufficiently distinct to warrant 
clear separation. 

The substitute heads of ankylosaurids differ 
from those of insects in two important respects. 
First, the substitute head of an insect is 
disposable: its sacrifice may permit an insect to 
escape with minor damage (Cott 1940; 
Swynnerton 1926; Carpenter 1941). By contrast, 
the tail club of an ankylosaurid was not 
detachable. Second, insects can escape by jumping 
or flying, whereas ankylosaurids were probably 
incapable of outrunning theropod dinosaurs 
(Coombs 1978; Thulborn 1982, 1990). 
Ankylosaurids were obliged to stand and fight, 
and for that reason their substitute head is 
necessarily adapted as a robust deterrent weapon. 

Insects possessing a substitute head often show 
appropriate peculiarities of behaviour (Cott 1940; 
Curio 1965, 1976; Wickler 1968). For instance, 
most butterflies rest head upwards on steep 
surfaces, but those with a substitute head 




FIGURE 4. Butterflies of the genus Thecla, each with 
'dummy head' (at left) bearing antenna-like filaments 
and eye-spot (adapted from Cott 1940; Wickler 1968). 



MIMICRY IN DINOSAURS 



155 



frequently rest upside down. Butterflies of the 
species Thecla togarna instantly turn through 1 80° 
on alighting, so that the substitute head points in 
the previous direction of flight; then, on the 
approach of a predator, the butterfly appears to 
take off in the 'wrong' direction. In some cases 
the substitute head bears an eye-spot and antenna- 
like filaments whose movements simulate those of 
real antennae (Fig. 4), and butterflies of the genus 
Deudoryx have been reported to walk backwards. 
Given these examples, it is conceivable that 
ankylosaurids equipped with a substitute head 
might also have indulged in some form of 
deceptive behaviour. 



Ankylosaurid defensive behaviour 

The foregoing analogies and constraints provide 
a framework for the following new model of 
ankylosaurid defensive behaviour. 

In normal circumstances ankylosaurids 
probably carried the tail club at ground-level; it is 
unlikely to have been carried so conspicuously as 
to attract the attention of predators. On the close 
approach of a predator, the tail may have been 
raised to become clearly visible (Carpenter 1984; 
Fig. 3a). If a predator moved in to attack, 
appropriate movements of the ankylosaurid tail 
could simulate those of a genuine neck and head. 
On luring the predator within reach of the tail 
club, the substitute head might be swung away as 
if it were attempting to escape. This apparent 
retreat of the prey's head could provoke the 
predator into lunging after it, whereupon the tail 
club would be swung back straight into the face of 
the predator. 

According to this model, ankylosaurids did not 
struggle to manouevre the tail club into a position 
suitable for striking an aggressor; instead, the 
aggressor was lured within reach of the tail club. 
Also, the tail club would strike at the aggressor's 
head, rather than its feet, thereby achieving the 
greatest deterrent effect. The model requires that 
an aggressor should respond to the two most 
important visual cues known to affect the 
behaviour of existing predators: movement (the 
apparent 'escape 1 of the prey's 'head') and shape 
(the sharp constriction behind the prey's 'head 1 ). 
Finally, a tail club disguised as a head would 
permit an ankylosaurid to deal with two or more 
predators, as each of them would fall prey to the 
same deception. 

This model finds similarities in the defensive 
adaptations of certain molluscs and beetles. Many 



eolid molluscs have vividly coloured dorsal 
papillae, which may be erected and waved about 
on the approach of an aggressor (Edmunds 1966, 
1974). The papillae contain nematocysts, as 
predators may discover to their cost. However, it 
is not clearly established that the papillae simulate 
items of prey, nor that their movements serve in 
distracting and luring predators; the papillae might 
equally well be interpreted as aposematic devices 
(Cott 1940). Similar uncertainties apply in the 
case of those carabid beetles that squirt an acid 
secretion from the tip of the tail. Although 
prominent white eye-spots may divert predators to 
the tail of the beetle (Marshall & Poulton 1902), 
these might, once more, be aposematic in function. 
In view of these uncertainties, it is impossible to 
identify any extant organism that duplicates the 
entire pattern of defensive behaviour proposed for 
ankylosaurids. Nevertheless, every component of 
that behavioural model has some counterpart 
among living organisms. For example, both 
snakes and cats are known to lure or distract other 
animals by twitching the tip of the tail (Wickler 
1968; Carpenter & Ferguson 1977). 



Mimicry? 

If the ankylosaurid tail club did function as a 
substitute head, it would probably qualify as an 
example of mimicry. Certainly, it would exemplify 
Wickler's concept of mimicry (1965, 1968), which 
required the deception of a signal-receiver (usually 
a predator). The more restrictive definition 
proposed by Vane-Wright (1976) placed no 
emphasis on deception, but depended instead on 
the outcome of interactions between the mimic, its 
model and the signal-receiver. Vane-Wright 
specifically excluded 'decoys and deflective 
marks 1 , including dummy heads, from the realm 
of mimicry. However, the ankylosaurid tail club 
may have been multifunctional - serving in turn 
as a deflective structure, a lure and a weapon. In 
performing the second of those functions, by 
luring an aggressor within reach, the tail club 
would merit inclusion in Vane-Wright's category 
II (synergic aggressive mimicry). The 
requirements of this category are: (1) that the 
mimic simulates an organism attractive to the 
signal-receiver, and (2) that on the signal- 
receiver's approach, the mimic interacts with it to 
the advantage of the model. In the case of 
ankylosaurids, the tail would simulate the 
generalized form of head and neck in other 
ornithischian dinosaurs (most obviously the 



156 



T. THULBORN 



ornithopods, Fig. 3b). Those other ornithischians 
would derive advantage, in Vane-Wright's words 
(1976: 36), 'through the removal or debilitation of 
their predators'. 

Such interpretation of the ankylosaurid tail club 
constitutes the first report of mimicry in dinosaurs. 
Although a tail club occurred in at least two 
genera of sauropod dinosaurs (Dong et al. 1989), 
this was comparatively small and might not have 
been functionally equivalent to that of 
ankylosaurids. Even so, the defensive model 
proposed for ankylosaurids might be extrapolated 
to certain stegosaurs, where conspicuous tail 
spikes (see Galton 1990) may have drawn the 
attention of predators before being employed as a 
deterrent weapon. 



Discussion 

Mimicry is a subject of enduring controversy 
among biologists. Wickler remarked (1968: 13) 
that 'one hundred years after Bates [1862] first 
clearly defined the concept of mimicry, a review of 
the literature listed 1 500 papers arguing for or 
against it. This amounts to roughly fifteen papers 
a year, or more than one a month.' Few examples 
of mimicry have been identified among vertebrate 
animals (Wickler 1968: 18), and not surprisingly 
this phenomenon is virtually unknown in the fossil 
record (Boucot 1990: 457). In these circumstances 
the foregoing hypothesis of mimicry in 
ankylosaurid dinosaurs might be regarded with 
some scepticism. It seems appropriate to examine 
two predictable objections to that hypothesis. 

First, it may be objected that hypotheses about 
the behaviour of extinct organisms are not 
amenable to scientific testing. In the present case 
it is difficult to imagine how the behavioural 
interactions of ankylosaurids and their predators 
could ever be corroborated or falsified. Although 
this objection is certainly valid, it does not 
necessarily condemn the proposed model of 
ankylosaurid defensive behaviour to the realm of 
unscientific speculation. That model meets the 
stringent requirements of a scientific hypothesis in 
two respects — congruence and productivity. 

With regard to congruence (or consilience), the 
model provides a single coherent explanation for 
all pertinent observations, particularly those 
concerning the anatomical peculiarities of 
ankylosaurids. By contrast, one conventional 
interpretation of ankylosaurid defensive behaviour 
involves a major inconsistency or internal 
contradiction: it requires that the ankylosaurid 



body plan should confer both great stability and a 
high degree of agility. To use a familiar analogy, 
that conventional interpretation requires that 
ankylosaurids should combine the stability of a 
four-wheel-drive vehicle with the turning circle 
and manoeuvrability of a motor-cycle. 
(Alternatively, one might try to envisage a single 
human possessing both the stability of a Sumo 
wrestler and the agility of a ballet dancer.) Such a 
combination of antagonistic physical properties 
may well be an unattainable ideal and it appears 
to be approached only in exceptional 
circumstances (e.g. in the case of a tank, or a 
bulldozer, with one caterpillar tread operating in 
reverse). Although there is a suggestion that 
stegosaurs were capable of fending off predators 
by pivoting very rapidly on the hindfeet alone 
(Bakker 1986), it seems much less likely that the 
broad-bodied ankylosaurids could have done so. 

The model proposed in this paper meets the 
requirement for productivity by generating 
predictions that are (in theory at least) 
scientifically testable. One such prediction, 
concerning the distribution of teeth-marks left by 
predatory dinosaurs, was mentioned earlier. Two 
more predictions, derived from the general 
principles of mimicry, are: (a) that the mimics 
(ankylosaurids) should have inhabited the same 
environments as their models (ornithopods), and 
(b) that the mimics should have been less 
abundant than their models (Wickler 1968: 46- 
48). Yet another prediction stems from the 
suggestion that a substitute head may originate 
through the elaboration of a duplicate head: 
consequently, one might expect the ancestors of 
ankylosaurids to have possessed a tail that was 
similar in size and overall shape to the head and 
neck combined. By contrast, conventional 
interpretations of ankylosaurid defensive 
behaviour seem to generate only a single 
prediction — namely, that the lower leg and ankle 
were those regions of the carnosaur body most 
likely to sustain injuries during encounters with 
ankylosaurids. The model proposed in this paper 
offers a conflicting prediction — that such injuries 
were more likely to affect the carnosaur skull. 

Second, it might be objected that the 
ankylosaurid tail bears only slight resemblance to 
the head and neck of an ornithopod, and that such 
a remote similarity would be unlikely to deceive a 
discriminating predator. This objection 
encapsulates the common belief that a successful 
mimic must show close and detailed resemblance 
to its model, on the assumption that some less 
exact resemblance to the model would be 



MIMICRY IN DINOSAURS 



57 



inadequate to deceive a signal-receiver. Similar 
assumptions were adopted by Punnet (1915) and 
Goldschmidt (1945), who maintained that 
mimicry could not arise through gradual processes 
of natural selection because the initial chance 
resemblance between mimic and model would be 
so slight as to confer no advantage. However, this 
assumption appears to be groundless: 
experimental studies with birds, insects and 
artificial models reveal that some mimics can, and 
do, derive advantage from only superficial 
similarity to their models (Duncan & Sheppard 
1965; Wickler 1968: 94; Edmunds 1974: 90-99). 
For instance, Brower et al. (1963: 80) stated 'as 
an experimentally demonstrated fact ... that even a 
remote resemblance between heliconiine 
butterflies can be advantageous 1 in conferring 
protection from predators. Moreover, it cannot be 
assumed that all predators are equally 
discriminating. Among birds, for example, the 
situation was summarized by Edmunds (1974: 92) 
as follows: 



it is likely that a whole spectrum exists from birds 
which recognize prey by a single visual cue, and are 
hence easily deceived by even a poor mimic, to birds 
which recognize prey by overall appearance or by 
many visual cues and which can distinguish almost 
perfect mimics from their models. 

It is possible that a similar spectrum existed 
among theropod dinosaurs, with some identifying 
their prey by means of its overall appearance 
(Gestalt perception) while others relied on one or 
more specific visual cues, such as shape and 
direction of movement. The less discriminating of 
those predatory dinosaurs might well have been 
deceived by ankylosaurids equipped with a 
dummy head. 



Acknowledgments 

I thank Kenneth Carpenter, Colin McHenry, 
Ralph Molnar and Tom Rich for their comments 
and discussions — sometimes sceptical, but always 
encouraging. 



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Nicolson: London. 



EMYDURA LAVARACKORUM, A NEW PLEISTOCENE TURTLE 

(PLEURODIRA : CHELIDAE) FROM FLUVIATILE DEPOSITS AT 

RIVERSLEIGH, NORTHWESTERN QUEENSLAND 

A. W. White and M.Archer 



Summary 

In 1986 the carapace, plastron and pelvic remains of a large chelid turtle were recovered from 
Pleistocene sediments on Riversleigh Station in northwestern Queensland. The fossil remains are 
described herein. The good state of preservation enabled the remains to be placed in an extant genus 
(Emydura) within the Chelidae on the basis of a derived feature of the carapace. Distinctive features 
such as wide bridge, broad first vertebral scute, unusual epiplastrahypoplastron suture, thickened 
bridge, buttresses and deep intergular insertion between the humerals prevent these fossils from 
being assigned to a known species. Accordingly, a new species of Emydura is proposed and 
comparisons are made between it and the currently recognised congenors. 



EMYDURA LAVARACKORUM, A NEW PLEISTOCENE TURTLE 
(PLEURODIRA:CHELIDAE) FROM FLUVIATILE DEPOSITS AT RIVERSLEIGH, 

NORTHWESTERN QUEENSLAND. 



A. W. WHITE AND M. ARCHER 



WHITE, A. W. & ARCHER, M. 1994. Emydura lavarackorum, a new Pleistocene turtle 
(Pleurodira: Chelidae) from fluviatile deposits at Riversleigh, northwestern Queensland. Rec. S. 
Aust. Mus. 27(2): 159-167. 

In 1986 the carapace, plastron and pelvic remains of a large chelid turtle were recovered from 
Pleistocene sediments on Riversleigh Station in northwestern Queensland. The fossil remains 
are described herein. The good state of preservation enabled the remains to be placed in an 
extant genus {Emydura) within the Chelidae on the basis of a derived feature of the carapace. 
Distinctive features such as a wide bridge, broad first vertebral scute, unusual epiplastra- 
hyoplastron suture, thickened bridge buttresses and deep intergular insertion between the 
humerals prevent these fossils from being assigned to a known species. Accordingly, a new 
species of Emydura is proposed and comparisons are made between it and the currently 
recognised congenors. 

Arthur W. White and Michael Archer, School of Biological Sciences, University of New South 
Wales, P.O. Box 1, Kensington, New South Wales, 2033. Manuscript received 17 September 
1993. 



Chelid fossil shell fragments are common in the 
Plio-Pleistocene deposits throughout Australia 
(e.g. in the Pliocene and Pleistocene deposits of 
the Darling Downs; Molnar 1982) but few are 
sufficiently complete to permit a detailed 
comparison with the shells of living species 
(Gaffney 1981). 

In May, 1984 exploration of eroded fluviatile 
deposits exposed in the watershed of the Gregory 
River on Riversleigh Station, northwestern 
Queensland, revealed several sites that produced 
fossil vertebrates. One of these, now known as 
Terrace Site, produced abundant remains of chelid 
turtles as well as the Pleistocene marsupial 
Diprotodon optatum, unidentified macropodoids, 
murids, crocodilians, lacertilians, fish and 
invertebrates (bivalves and gastropods). 
Preliminary accounts of the discovery of Terrace 
Site have been given in Archer, Hand and 
Godthelp(1986). 

From approximately 30cm above the base of the 
palaeochannel, in a layer with abundant bivalves 
and mammal bone fragments, a nearly complete 
although somewhat crushed chelid shell was 
recovered. It is sufficiently well preserved to 
enable comparison with a wide range of living 
chelids and is the basis for the species described 
herein. 

Additional vertebrate remains from this deposit 
will be reported elsewhere (Godthelp, in 



preparation; Willis and Archer 1990; Willis, in 
preparation). 

The anatomical terminology used in this paper 
follows that employed by (Gaffney 1977). That of 
modern chelid taxonomy follows Cogger (1993) 
and Obst (1986). Qualitative and quantitative 
methods of analysis follow Auffenberg (1976). 



Methods 

After being exposed in the deposit, the specimen 
was hardened with aquadhere, braced and secured 
in a plaster jacket. In the laboratory, various 
sections of the shell, defined by post-depositional 
crushing, were reconstructed using aquadhere. 
Measurements were made with dial calipers to the 
nearest millimetre. 

Measurements and abbreviations used here are as 
follows: 



Carapace 

Anterior Carapace Width (ACW) = the anterior 
width of the carapace measured from the most 
anterior points of the M 2 - M 3 sutures. 

Width of V f (V W) = the maximum width of the 
first vertebral scute. 



160 



A. W. WHITE & M. ARCHER 



Length ofVj (V } L) 
first vertebral scute. 



the maximum length of the 



Width and Length of Subsequent Scutes (Indicated 
by the appropriate subscript). 

Length of C } (C L) = the maximum length of the 
first costal scute. 



Entoplastron Length (EnL) = the maximum length 
of the entoplastron. 

Entoplastron Width (EnW) = the maximum width 
of the entoplastron. 

Hypoplastron Length (HyL) = the length of the 
hypoplastron bones along the midline. 



Plastron 

Total Plastron Length (TPL) = measured in 
parallel to the midline from the most anterior to 
most posterior part of the plastron. 

Anterior Plastron Length (APL) = measured in 
parallel to the midline from the most anterior 
portion of the plastron to the anterior margin of 
the bridge. 

Bridge Width (BW) = the width of the bridge at 
the junction of the plastron. 

Posterior Plastron Length (PPL) = measured in 
parallel to the midline from the most posterior 
portion of the plastron to the posterior margin of 
the bridge. 

Intergular Width (IW) = the width of the intergular 
measured along the anterior margin of the 
plastron. 

Intergular Length (IL) = the maximum length of 
the intergular scute. 

Gular Width (GW) = the width of the gulars along 
the anterior margin of the plastron. 

Gular Length (GL) = length of the gular-intergular 
suture. 

Intergular Insertion (II) = the distance along the 
midline that the intergular scute penetrates 
between the humeral scutes i.e. measured from a 
line level with the posterior ends of the gulars to 
the posterior end of the intergular. 

Humeral Length (HL) = the length of the humeral 
scutes along the midline. 

Pectoral Length (PL) - the length of the pectoral 
scutes along the midline. 

Abdominal Length (AL) - the length of the 
abdominal scutes along the midline. 

Femoral Length (FL) = the length of the femoral 
scutes along the midline. 

Anal Length (AnL) - the length of the anal scutes 
along the midline. 

Anal Width (AW) = the distance between the most 
posterior parts of the opposing anal scutes. 

Epiplastron Length (EpL) — the length of the 
epiplastron bones along the midline. 



Comparative Specimens Examined 

Many of the comparative specimens of modern 
species examined during this study are lodged in 
the herpetological collections of the Australian 
Museum as follows: 

Emydura australis: R20737, R72786, R72787. 

Emydura kreffti: R14925. 

Emydura macquarii: R1188, R6789, R81477, 
R85727, R104335,R123049 

Emydura novae guineae: R5042, R24460. 

Emydura signata: R58588, R58589, R96716. 

Elseya dentata: R3699, R3700, R31728, R36998, 
R40181, 

Elseya latisternum: R20330-20345,R21224, 
R21485, R21570-21572, R37657-37665, 
R43530, R43542, R81958. 

Rheodytes:R\ 25481 

Other specimens of these species were 
examined from the authors' collections. A 
specimen of Pseudemydura umbrina was 
examined from the Western Australian museum 
(WAM 13744). A specimen of the alpha taxon 
turtle was examined from John Cann's collection. 



Systematics 

Order: Testudines Linneus, 1758 
Infraorder: Pleurodira (Cope, 1 868) 
Family: CHELIDAE Gray, 1825 

Genus: Emydura Bonaparte, 1836 

Emydura lavarackorum White & Archer n. sp. 
(Figs 1,2,3 and 4) 

Holotype: 

Queensland Museum Palaeontological 
Collections no. F 24121, an associated almost 
complete plastron, partial carapace and pelvic 
fragments collected 9th May, 1986, by J. and S. 
Lavarack. 



NEW PLEISTOCENE TURTLE 



161 



Type locality and age: 

Terrace Site, an excavation in fluviatile 
sediments exposed on the south bank of the 
Gregory River, Riversleigh Station, northwestern 
Queensland, approximately 200km northwest of 
Mount Isa. More precise locality data are recorded 
and may be available on application to the 
Queensland Museum or the University of New 
South Wales. The presence in the sediments of 
material referable to Diprotodon optatum and no 



other index fossil indicative of any other period of 
time is the basis for interpreting the deposit to be 
Pleistocene in age. No more precise age 
determination is available at this time although 
samples suitable for radiocarbon dating have been 
collected. 

Diagnosis: 

This species differs from all others in the 
following combination of features. The first 




10 cm 



FIGURE 1 . Dorsal view of anterior carapace of Emydura lavarackorum. 



162 



A. W. WHITE & M. ARCHER 



vertebral scute (V { ) is much wider than V , 
Vertebral scutes V 2 and V 3 are rectangular, being 
longer than they are broad, with very small 
projections into the costal junctions (Fig.l). The 
humeral-pectoral seam is sigmoidal rather than 
straight. The anterior straight edge of the carapace 
is wide and incorporates the two left and right 
marginal scutes (before the carapace curves 
posteriorly). The anterior bridge struts are 
unusually thick. The anterior edge of the gular is 
as wide as the anterior edge of the intergular. The 
intergular is long and deeply divides the humeral 




1 o cm 



FIGURE 2. External view of plastron of Emydura 
lavarackomm 



scutes. The intergular intrusion between the 
humerals is greater than the gular length. The 
intergular scutes narrow at the anterior edge of the 
carapace. The bridge is broad (BL:TPL = 0.29). 
The acetabulum is circular and is contributed to 
equally by all three pelvic bones. The acetabulum 
has a diameter of 25 mm. The upper (ilial) lip of 
the acetabulum is raised and overhanging whereas 
the lower lip (ischium and pubis) is less 
pronounced. 

Etymology: 

The species name is in honour of Sue and Jim 
Lavarack, hard-working volunteers who, besides 
having collected the holotype (Archer 1988) and 
supervised excavations at Terrace Site for five 
years, have maintained a continuous supportive 
role in the work done at Riversleigh and on 
Riversleigh materials which they have helped to 
prepare in Sydney. 

Description: 

The plastron is long (390 mm) and almost 
complete except for some medial gaps in the anal 
region (Fig. 2). The plastron is evenly rounded at 
its anterior end. The posterior end of the plastron 
terminates with two pointed anal projections. The 
anterior lobe of the plastron is broader (maximum 
width 165 mm) than the posterior lobe (maximum 
width 154 mm). 

The endoplastron is wider (EnW = 58 mm) than 
it is long (EnL = 45 mm) (Fig. 3). The epiplastral- 
hypoplastral suture is sigmoidal. The hypoplastra 
are the longest bony elements in the plastron (HyL 
= 105 mm). Of the epidermal scutes, the intergular 
is the most distinctive. It completely separates the 
gulars and penetrates deeply between the 
humerals. The humeral-humeral seam is only 15 
mm long whereas the humerals have a maximum 
height of 87 mm. The longest scutes are the ab- 
dominal (A, = 105 mm), followed by the femoral 
scutes (F = 95 mm), pectoral scutes (PL = 67 
mm), anal scutes (AnL = 66 mm), intergular (IL = 
62mm) and finally the humerals (HL= 15 mm). 

The humeral-pectoral seam is sigmoidal, the 
most anterior sections being at the midline and at 
the extreme margins. The intergular has a 
maximum width of 26 mm but is only 19 mm 
wide along the anterior edge of the plastron. The 
gulars are small being only a little wider (GL = 28 
mm) than the intergular. The intergular extends 
most of the way between the humerals. The 
intergular intrusion is longer than the gular length. 

The carapace is large and flat along the ventral 
surface. The leading edge of the carapace is almost 



NEW PLEISTOCENE TURTLE 



163 




FIGURE 3. Internal view of plastron of Emydura 
lavarackorum. 





straight and does not curve posteriorly until the 
suture line between the second and third marginal 
scutes. A precentral (nuchal) scute is absent. The 
broad anterior edge of the carapace is reflected by 
an expansion of the V, scute (maximum width 
103 mm). This scute is almost twice as wide as 
the second vertebral scute (width 56 mm). V t 
(height 70 mm) is not as high as V 2 (height 87 
mm). The third vertebral scute is incomplete but 
appears to be of similar proportions to the V 2 
scute. The projections of the vertebral scutes 
between the costals is minimal. The first costal 
scute (Cj) is higher (95 mm) than C 2 (83 mm in 
height). It was not possible to measure the width 
of the costals. 

The recess for the insertion of the anterior 
bridge strut on the undersurface of the carapace is 
steeply angled across the first pleural bone and is 
not in line with the raised process that forms the 
mid-pleural wall. The recess abuts the second 
peripheral bone and forms most of the base of the 
third peripheral (Fig 5). The dorsal fork of the 
transverse process on the first thoracic vertebra 
sweeps backwards to form the top of the mid- 
pleural wall. 

A major section of the left pelvis comprising of 
the ischium, ilium and pubis was measured. The 
piece was 55 mm long and 20 mm wide at the 
ilial fracture. It was 35 mm wide at a position 
level with the acetabulum (which had a diameter 
of 25 mm). 

The acetabulum is quite deep and is overhung 
by a pronounced upper ridge formed by the 
extension of the ilial ridge (Fig 4). The lower lip 
of the acetabulum has a much weaker rim formed 
a joining ridge continuous between the pubis and 
ischium. 



FIGURE 4. Acetabular view and lateral view of the left hip 
of Emydura lavarackorum. 



Measurements (mm) of the holotype: 

ACW = 210; V,W =105; V,H = 72; V 2 W = 65; 
V 2 H = 87; C,H = 95; C 2 H = 83; TPL = 390; APL 
= 110;BW= 115; PPL = 165; IW = 20; IH = 62; 
GW = 30; GL = 30; II = 36; HL = 15; PL = 67; 
AL = 105; FL = 95; AnL = 66; AW = 93; EpL = 
30; EnH = 45; EnW = 58; HyL = 105. 

Comparisons and discussion 

The Riversleigh fossil turtle is placed in the 
Chelidae because of the evidence of pelvic fusion 
to the shell and the absence of mesoplastra and 
neural bones (Gaffney 1977). 

Within the Chelidae, the Riversleigh turtle is 
placed in the genus Emydura on the basis of a 
recently identified synapomorphy. This feature 
relates to the insertion of the anterior bridge into 



164 



A. W. WHITE & M. ARCHER 



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NEW PLEISTOCENE TURTLE 



165 



the ventral surface of the carapace. The recess for 
the anterior bridge is angled steeply backwards to 
reach the raised process that forms a transverse 
wall across the floor of the first pleural bone. In 
the other chelid genera, the recess is itself 
transverse and so forms a near continuous line 
with the mid-pleural wall (Fig. 5). The Riversleigh 
Emydura has this derived characteristic. 

Osteological comparisons (e.g. Gaffney 1977) 
of the shells and skulls of Emydura and Elseya 
have highlighted the great degree of similarity 
between these two genera. Current taxonomy 
(Obst 1986, Cogger 1993) recognises two species 
of Elseya and 6 species of Emydura. The 
diagnostic features that are used to identify the 
genera and species are features that include soft 
anatomy, skull ridges and plastral scute patterns. 
Legler (1985) and Georges and Adams (1993) 
have cast doubts about the validity of current 
concepts of these genera, especially Elseya which 
appears to be paraphyletic. On the basis of the 
carapacial synapomorphy identified in this paper 
Emydura, however, appears to be monophyletic. 

The Riversleigh Emydura is a large turtle 
compared to other Australian chelids. It has a 
plastral length of 39 cm and hence would most 
likely have had a carapace length of approximately 
41 cm. In comparison, the largest measured extant 
chelid is Elseya dentata. Cann (1986) has reported 
adults of this species with carapace lengths of up 
to 36.5 cm. The largest Emydura species is E. 
macquarii which may reach shell lengths of up to 
40 cm (Cogger 1993). 

In the extant Australian chelid species, the 
bridge typically occupies between 20 and 30% of 
the plastron length (Table 1). Emydura macquarii 
and E. signata have the narrowest bridge (20- 
23% of the plastron length) of all Emydura 



species. In contrast the bridge of E. kreffti is 
particularly broad and varies between 28 and 33% 
of the plastron length In the Riversleigh turtle, the 
bridge is wide (30% of the plastron length). 

The bridge struts of the Riversleigh turtle are 
unusually thick. The anterior bridge struts are 2.15 
cm wide at the base and almost 1cm wide in the 
middle. The largest comparative chelid specimen 
that was available for measurement was a deep- 
shelled snapper (E. dentata carapace length 28.5 
cm, R40181). The anterior bridge strut of this 
animal was only half as massive. 

Carapace shape varies in Emydura. Species 
such as E. macquarii and E. signata have broad, 
low domed carapaces that are expanded at the 
rear. In the range of shell shapes E. kreffti 
represents the other extreme and has a high domed 
carapace that is not expanded at the rear (Goode 
1966). E. lavarackorum is intermediate in shell 
shape and has a carapace that is not evenly 
rounded. 

In all species of Emydura the gular scutes of the 
plastron are completely divided by the intergular 
(Table 1). Emydura turtles also show partial 
separation of the humerals by the intergular, with 
the degree of separation being less than half of the 
mid-humeral length. Of the other Australian 
chelid genera, only in Pseudemydura does the 
intergular also completely divide the humerals. 
The intergular of Elseya latisternum produces the 
weakest separation of the humerals and barely 
intrudes between the humerals. Emydura 
lavaracki is most distinctive in this regard as the 
intergular deeply divides the humerals without 
completely separating them. 

In the extant Australian Emydura, the 
arrangement of the pelvic bones around the 
acetabulum is similar. Here the ilium, ischium and 





FIGURE 5. View of the undersurface of the anterior carapaces of Emydura kreffti (a) and Elseya latisternum (b). 



166 



A. W. WHITE & M. ARCHER 



pubis contribute almost equally to the composition 
of the acetabulum. This arrangement is also 
present in Emydura lavarackorum. In the extant 
species of Emydura the upper and lower rim of 
the acetabulum are equally protuberant. E. 
lavaracki is different in this regard as the upper 
lip of the acetabulum is more substantial than the 
lower lip. This means that the acetabulum is 
relatively deeper in this species. 

The conventional guide for determining species 
within the genus Emydura is based on features of 
soft anatomy, shell shape and distribution (Cogger 
1993). Other shell features have been used such as 
the relative length of the plastron compared to the 
carapace, the relative length of the anterior 
plastron compared to the posterior plastron and 
the relative width of the bridge (Goode 1966). E. 
macquarii has a plastron that is noticeably shorter 
than the carapace (80-85% of the carapace 
length). By way of contrast, E. kreffti has a longer 
plastron that ranges from 85-95% of the carapace 
length. Based on the shell reconstruction of E. 
lavarackorum the plastron appears to be about 
95% of the length of the carapace. In some species 
of Emydura, such as E. macquarii, the anterior 
plastron is conspicuously shorter than the posterior 
plastron. In comparison, E. kreffti has anterior and 
posterior plastron segments that are almost equal 
in length. E. lavarackorum the anterior plastron is 
much shorter (67%) than the posterior plastron. 

In view of the many evidently significant 
differences between the Riversleigh fossil species 
and any others referred to this genus, we have no 
hesitation in describing the fossil form as the new 
species Emydura lavarackorum. 

Although we have placed this species in the 
genus Emydura on the basis of shell morphology 
it is apparent that it is not particularly closely 
related to any of the other species in this genus. In 
two respects, this Pleistocene chelid is unlike all 



Australian short-necked turtles: the massive 
expansion of the W l scute and the deep extension 
of the intergular between the numerals. In these 
features, the Riversleigh form more closely 
resembles the long-necked chelids. Further 
clarification of the intrafamilial affinities of the 
Riversleigh form may have to await discovery of 
cranial material. 



Acknowledgments 

We wish to acknowledge the vital financial support 
the Riversleigh Project has had from: the Australian 
Research Grant Scheme (Grant PG A3 851506P); the 
National Estates Scheme (Queensland); the Department 
of Environment and Arts; Wang Australia Pty Ltd; ICI 
Australia Pty Ltd; the Queensland Museum; the 
Australian Geographic Society; Mount Isa Mines Pty 
Ltd; and Surrey Beatty & Sons Pty Limited. Critical 
logistical support in the field and laboratory has been 
received from the Riversleigh Society, the Friends of 
Riversleigh, the Royal Australian Air Force, the 
Australian Defence Force, the Queensland National 
Parks and Wildlife Service, the Riversleigh Consortium 
(Riversleigh being a privately owned station), the Mount 
Isa Shire, the Burke Shire, the Northwest Queensland 
Tourism and Development Board, the Gulf Local 
Development Association, PROBE and many volunteer 
workers and colleagues including Dr Suzanne Hand and 
Henk Godthelp who helped to recover the holotype. Alan 
Rackham Snr and Alan Rackham Jnr discovered the 
Terrace Site in the first place. Among the volunteers who 
helped at the 1984 Riversleigh Expedition, special thanks 
are extended to Bruce Armitage, John Courtenay, Martin 
Dickson, Joy Dohle, Archie Daniels, Bernie Jennings, 
Leslie Hobbs, Bill Lockwood, Audrie and Harry 
Macintosh, Reg Markham, Melody Nixon, Ron Norman, 
Jim and Sue Lavarack, Brian, Bronwen and Dan 
O'Dwyer, Lazar Radanovich, Jim Ross, Mary 
Sandilands, Donald and Sue Scott-Orr, Jack Struber, Dr 
Steven Waite and Shirley Webster. 



References 



AUFFENBERG, W. 1976. The genus Gopherus 
(Testudinidae) Part 1 . Osteology and relationships of 
extant species. Bulletin of the Florida State Museum 
of Biological Sciences 20: 47-1 10 

ARCHER, M., HAND, S. H. & GODTHELP, H. 1986. 
'Uncovering Australia's Dreamtime'. Surrey Beatty & 
Sons Pty Ltd: Sydney. 

ARCHER, M. 1988. Riversleigh: window into our 
ancient past. Australian Geographic 9: 40-57 
[illustration of the holotype of Emydura 
lavarackorum being collected at Terrace Site by J. 
Lavarack, p. 53]. 



CANN, J. 1978. 'Tortoises of Australia'. Angus & 
Robertson, Sydney. 

COGGER, H. G. 1993. 'Reptiles and Amphibians of 
Australia'. 5th ed. Reed, Sydney. 

COGGER, H. G., CAMERON, E. E. & COGGER, H. 
M. 1983. Amphibia and Reptilia. In 'Zoological 
Catalogue of Australia'. Vol. 1. Ed. D. W. Walton. 
Bureau of Flora and Fauna: Canberra. 

GAFFNEY, E. S. 1977. The short-necked family 
Chelidae: a theory of relationships using shared 
derived characters. American Museum Novitates 
2620: 1-28. 



NEW PLEISTOCENE TURTLE 



167 



GAFFNEY, E. S. 1981. A review of the fossil turtles of 
Australia. American Museum Novitates 2720: 1-38. 

GEORGES, A. & ADAMS, M. 1993. A phylogeny for 
Australian chelid turtles based on allozyme 
electrophoresis. Australian Journal of Zoology 40: 
453-476. 

GOODE, J. 1966. 'Freshwater Tortoises of Australia and 
New Guinea'. Longman Press. 

LEGLER, J. M. 1985. Australian chelid turtles: 
Reproductive patterns in wide ranging taxa. Pp. 117- 
123 in 'Biology of Australian Frogs and Reptiles'. 
Eds G. C. Grigg, R. Shine and H. Ehmann. Royal 
Zoological Society of N.S.W. Surrey Beatty & Sons. 

LEGLER, J. M. & CANN, J. 1980. A new genus and 



species of Chelid turtle from Queensland, Australia. 
Contributions of the Science and Natural History 
Museum of Los Angeles County 324: 1-18. 

MOLNAR, R. 1982. Cenozoic Fossil Reptiles in 
Australia. Pp. 228-233 in 'The Fossil Vertebrate 
Record of Australasia'. Eds P. V. Rich and E. M. 
Thompson, Monash University Press, Victoria. 

OBST, F. J. 1986. 'Turtles, Tortoises and Terrapins'. 
Druckerei Fortschritt Erfut, Leipzig, G.D.R. 

WILLIS, P. & ARCHER, M. 1990. A Pleistocene 
longirostrine crocodile from Riversleigh; first 
occurrence of Crocodilus johnstoni Krefft. Memoirs 
of the Queensland Museum 28: 159-164. 



FROGS FROM A PLIO-PLEISTOCENE SITE AT FLORA VILLE STATION, 

NORTHWEST QUEENSLAND 

Michael 7. Tyler, Henk Godthelp & Michael Archer 



Summary 

Six frog ilia, mostly in good condition, have been recovered from riverine granuals on the western 
bank of the Leichhardt River, south of the Floraville Station in northwest Queensland. 



FROGS FROM A PLIO-PLEISTOCENE SITE AT FLORA VILLE STATION, 

NORTHWEST QUEENSLAND 



MICHAEL J. TYLER, HENK GODTHELP & MICHAEL ARCHER 



TYLER, M. J., GODTHELP, H. & ARCHER, M. 1994. Frogs from a Plio-Pleistocene site at 
Floraville Station, northwest Queensland. Rec, S. Aust. Mus. 27(2): 169-173. 

Six frog ilia, mostly in good condition, have been recovered from riverine gravels on the 
western bank of the Leichhardt River, south of the Floraville Station Homestead at Floraville 
Station in northwest Queensland. 

Three species of frogs are included: the extant species Cyclorana cultripes Parker, C. 
platycephala (Gunther) and also Limnodynastes sp. cf. L. tasmaniensis Giinther. They 
constitute the first fossil record of the fossorial genus Cyclorana, whilst the Limnodynastes 
species resembles closely material from earlier (Oligo-Miocene) sites at the nearby locality of 
Riversleigh Station. 

M. J. Tyler, Department of Zoology, University of Adelaide, South Australia 5005, and H. 
Godthelp and M. Archer, Vertebrate Palaeontology Laboratory, School of Biological Sciences, 
University of New South Wales, Kensington, New South Wales 2006. Manuscript received 12 
May 1993. 



A characteristic of the frog fossil record in 
Australia is that most of the Tertiary sites are in 
the north of the continent, whereas the Quaternary 
sites are predominantly in the south (Tyler, 1989, 
in press; Tyler & Godthelp, 1993). Here we report 
an exception to the trend, for one of us (M. A.) 
recovered vertebrate material from an undated site 
in northwest Queensland considered to be Plio- 
Pleistocene. The site was named 5C in the M. A. 
notebook and consists of riverine gravels located 
on the western bank of the Leichhardt River south 
of the Floraville Station Homestead. Included are 
six anuran ilia representing three species. 



Materials and methods 

The ilia are deposited in the collection of the 
Queensland Museum, Brisbane. Methods of 
measurement and descriptive terminology follow 
Tyler (1976). Scanning electron micrographs were 
prepared with a Cambridge Autoscan Model 
5250. 



System atics 



Family HYLIDAE 

Sub-family PELODRYADLNAE 

Genus Cyclorana Steindachner 



The generic features of the pelvis of Cyclorana 
were described by Tyler (1976) based on the 
examination of the closely related taxa C. 
australis (Gray) and C novaehollandiae 
Steindachner, and the more divergent C. dahlii 
(Boulenger) and C. platycephala (Gunther). The 
only major difference in ilial structure noted 
amongst these species was the absence in C. 
platycephala of a narrow dorsal rim possessed by 
the others. 

Subsequently C dahlii was transferred from 
Cyclorana to the genus Litoria Tschudi by Tyler, 
Davies & King (1978). For that reason and 
because of access to all Cyclorana species except 
C manya, it now is possible to redefine the ilial 
characteristics of Cyclorana. 

The ilial shaft is long, slender and slightly 
curved, and in C. australis and C. novae- 
hollandiae bears a narrow dorsal rim on the lateral 
surface, and a corresponding depression upon the 
medial surface. This rim and indentation is 
lacking in the remaining species. The acetabular 
rim is narrow and the fossa extensive. 

The ventral acetabular expansion is gently 
rounded and the preacetabular zone is narrow. The 
dorsal acetabular expansion is well developed and 
has a gently curved anterior margin. There is a 
major dichotomy in the form of the dorsal 
protuberance: inconspicuous and laterally 
disposed in C. australis and C. novaehollandiae, 
but distinctly elevated as a very conspicuous 
feature in the remaining congeners. 



170 



M. J. TYLER, H. GODTHELP & M. ARCHER 




FIGURES 1-3. Cyclorana cultripes Parker. I: QM F 23023; 2: QM F 23024; 3: QM F 23029. 



PLIO-PLEISTOCENE FROGS 



171 




FIGURES 4-6. Cydorana and Limnodynastes. 4: Cyclorana platycephala (Giinther), QM F 23210; 5: Cyclorana 
platycephala (Giinther), QM F 23025; 6: Limnodynastes sp. cf. L. tasmaniensis Giinther, QM F 23022. 



172 



M. J. TYLER, H. GODTHELP & M. ARCHER 



Cyclorana cultripes Parker 
Figs 1-3 

Material: QM F 23023, left ilium; F 23024 left 
ilium; F 23029 left ilium. 

The ilial characteristics of C. cultripes have not 
been reported. The fossil material listed above 
conforms to ilia dissected from extant material 
principally in the unusual form of the dorsal 
prominence on the ilial shaft which is elongate, 
and extends medially over much of its length. As 
indicated in Figs 1-3 the dorsal prominence has a 
length equivalent to the diameter of the ilial 
portion of the acetabular fossa. In each of the 
fossils the lateral extremity of the dorsal 
prominence has been abraded. 



Cyclorana platycephala (Gunther) 
Figs 4-5 

Material: QM F 23210, left ilium; F 23025 right 
ilium. 

In its overall habitus C. platycephala is the 
most divergent of all of the members of this 
genus. What sets it apart is the large flat head 
with small eyes protuding from the dorsal surface, 
and the extensively (usually fully) webbed toes. 
The ilium is equally distinctive by virtue of the 
highly developed dorsal prominence, which rises 
high above the ilial shaft in an almost cylindrical 
form. 



Family LEPTODACTYLIDAE 
Sub-family LIMNODYNASTINAE 



"A) 


A, 




AUSTRALIA 



Major Roads / 



00 



N 



km 



16- 


B 


Gulf of 
_^ Carpentaria 

(7 / 


i 




\° 1 


18- 




; 

• Floraville 




1 Riversleigh ; 
. Station 




•- 


'*;, >'- N 


20°- 


w-,. 






1 


\ Mt Isa \ 

i 



138 



138 



FIGURE 7. Location of Floraville Station and Riversleigh 
Station, Queensland. 



Genus Limnodynastes Fitzinger 



Limnodynastes sp. cf, L, tasmaniensis Gunther 
Fig. 6 

Material: QM F 23022, left ilium. 

Amongst the members of the genus 
Limnodynastes the features that are unique to this 
species are the protuberant nature of the sub- 
acetabular zone, the lateral groove upon the ilial 
shaft, and the anteriorly inclined dorsal 
prominence and dorsal protuberance. These 
features are shared by modern representatives of 
Limnodynastes tasmaniensis, and particularly 
specimens reported from Oligo-Miocene sites at 
Riversleigh Station in northwest Queensland 



(Tyler, 1990a). The location of Floraville and 
Riversleigh Stations is shown in Fig. 7. 



Discussion 

This small assemblage of frogs from the 
Tertiary of northwest Queensland is significant for 
several reasons. Firstly because of the presence of 
two extant species of Cyclorana, being the first 
fossil record of this genus. Neither species has 
been reported as far northeast. In the case of C. 
platycephala every voucher specimen known 
throughout Australia until January 1990 was 
plotted by Tyler (1990b). Floraville Station is just 
beyond its known modern geographic range. The 
presence of C. cultripes at Floraville Station 



PLIO-PLEISTOCENE FROGS 



173 



similarly has not been recorded, but it is common 
at similar latitudes and may be assumed possibly 
to exist there today; there has been minimal 
collecting activity in the area. 

The presence of Limnodynastes sp. cf. L. 
tasmaniensis in this collection is intriguing. The 
modern species has an extensive distribution 
throughout eastern and southeastern Australia, and 
was introduced to Kununurra in the Kimberley 
Division of northern Western Australia (Martin & 
Tyler 1978). More recently a second isolated 
population has been located at Newry Station in 
the Northern Territory (Tyler, Watson, & Davies, 
1983). The interpretation of the presence of the 
species so far from its known geographic range is 
that it was introduced there. The location of 
comparable material at mid-Miocene sites at 
Riversleigh Station in northwest Queensland 
(Tyler 1990a), and its presence at Floraville 
Station in the Plio-Pleistocene collectively suggest 
a far longer duration of existence in northern 
Australia of an ancestral form resembling the 



modern. Clearly it is possible that the extant 
Kununurra (WA) and Newry Station (NT) 
populations should be explored as possible relics, 
rather than recent introductions. 

The mammal fauna at the site is dominated by 
several taxa of as yet undescribed rodents. There 
is also a high diversity of marsupials, with some 
extant genera. The sediments are undated but are 
suggested to be Plio-Pleistocene by Archer (1982, 
1984). 



Acknowledgments 



We are grateful to Mr and Mrs Camp for permission 
for M. A. to work on Floraville Station. The field work 
was supported by R. E. Lemley, the Queensland 
Museum and the Australian Research Grants Committee. 
Comparative studies were aided by S. Bryars and 
supported by the Australian Research Council. The 
S.E.M. photographs were prepared by Stuart Mclure of 
the CSIRO Division of Soils, Adelaide. 



References 



ARCHER, M. 1982. Review of the dasyurid 
(Marsupialia) fossil record, integration of data bearing 
on phylogenetic interpretation, and suprageneric clas- 
sification. Pp. 595-619 in 'Carnivorous marsupials' 
vol. 2. Ed. M. Archer. Royal Zoological Society of 
New South Wales. 

ARCHER, M. 1984. The Australian marsupial radiation. 
Pp. 633-808 in 'Vertebrate Zoogeography and Evolu- 
tion in Australasia'. Ed. M. Archer and G. Clayton. 
Hesperian Press. 

MARTIN, A. A., & TYLER, M. J. 1978. The introduc- 
tion into Western Australia of the frog Limnodynastes 
tasmaniensis. Australian Zoologist 19 (3): 321-325. 

TYLER, M. J. 1976. Comparative osteology of the pelvic 
girdle of Australian frogs and description of a new 
fossil genus. Transactions of the Royal Society of 
South Australia 100(1): 3-14. 

TYLER, M. J. 1989. 'Australian Frogs'. Viking O'Neil, 
Melbourne. 

TYLER, M. J. 1990a. Limnodynastes Fitzinger (Anura: 
Leptodactylidae) from the Cainozoic of Queensland. 
Memoirs of the Queensland Museum 28(2): 779-784. 



TYLER, M. J. 1990b. Geographic distribution of the 
fossorial hylidfrog Cyclorana platycephala (Gunther) 
and the taxonomic status of C. slevini Loveridge. 
Transactions of the Royal Society of South Australia 
114(2): 81-85. 

TYLER, M. J. in press. Hylid frogs from the Mid-Mio- 
cene Camfield Beds of Northern Australia. Beagle. 

TYLER, M. J., DAVIES, M. & KING, M. 1978. The 
Australian frog Chiroleptes dahlii Boulenger: its sys- 
tematic position, morphology, chromosomes and dis- 
tribution. Transactions of the Royal Society of South 
Australia 102(1): 17-24. 

TYLER, M. J., WATSON, G. F. & DAVIES, M. 1983. 
Additions to the frog fauna of the Northern Territory. 

Transactions of the Royal Society of South Australia 
107(4): 243-245. 

TYLER, M. J. & GODTHELP, H. 1993. A new species 
of Lechriodus Boulenger (Anura: Leptodactylidae) 
from the early Eocene of Queensland. Transactions of 
the Royal Society of South Australia 117(4): 187— 
189. 



FORM AND FUNCTION IN SCALES OF LIGULALEPSIS TOOMBSI 
SCHULTZE, A PALAEONISCOID FROM THE EARLY DEVONIAN OF 

AUSTRALIA 

Carole Burrow 



Summary 

A variety of isolated scales and lepidotrichia from the Early Devonian of New South Wales is 
referred to the palaeoniscoid Ligulalepis toombsi Schultze 1968, to date known only by isolated 
scales. Scales of various forms are attributed to specific regions of the body, on the basis of 
comparisons with articulated Palaeozoic palaeoniscoids. Mobility of scales in the forward flank 
region appears to have been constrained by a prominent process projecting from the rostrodorsal 
corner, in addition to the peg, socket and keel of typical palaeoniscoid scales. Such complex 
interlocking of the scales implies that the body of Ligulaepis toombsi was relatively inflexible. 
Some suggestions are offered regarding the possible mode of locomotion. 



FORM AND FUNCTION IN SCALES OF LIGULALEPIS TOOMBSI SCHULTZE, 
A PALAEONISCOID FROM THE EARLY DEVONIAN OF AUSTRALIA 



CAROLE BURROW 



BURROW, C. 1994. Form and function in scales of Ligulalepis toombsi Schultze, a 
palaeoniscoid from the early Devonian of Australia. Rec. S. Aust. Mus. 27(2): 175-185. 

A variety of isolated scales and lepidotrichia from the Early Devonian of New South Wales is 
referred to the palaeoniscoid Ligulalepis toombsi Schultze 1968, to date known only by isolated 
scales. Scales of various forms are attributed to specific regions of the body, on the basis of 
comparisons with articulated Palaeozoic palaeoniscoids. Mobility of scales in the forward flank 
region appears to have been constrained by a prominent process projecting from the rostro- 
dorsal corner, in addition to the peg, socket and keel of typical palaeoniscoid scales. Such 
complex interlocking of the scales implies that the body of Ligulalepis toombsi was relatively 
inflexible. Some suggestions are offered regarding the possible mode of locomotion. 

This is a contribution to IGCP 328: Palaeozoic Microvertebrates. 

Carole Burrow, Department of Zoology, University of Queensland, St Lucia 4072 Australia. 
Manuscript received 21 April 1993. 



Ligulalepis toombsi Schultze 1968 is the only 
palaeoniscoid so far described from Australian 
sediments older than the Late Devonian. Although 
no whole, or even partially articulated, specimens 
have yet been discovered, Ligulalepis scales are 
distinctive and widespread in microvertebrate 
assemblages from marine Lower Devonian 
deposits from south eastern, and possibly western, 
Australia. 

Scales of Ligulalepis toombsi were first 
described by Schultze (1968) from the lower part 
of the Murrumbidgee Group of Taemas, NSW, in 
greatest abundance from the Spinella yassensis 
Limestone Member of early Emsian age (probable 
conodont zone dehiscens or gronbergi - Campbell 
and Barwick 1988, Talent 1989). Ligulalepis 
toombsi scales were also reported by Giffin (1980) 
in a diverse microvertebrate assemblage from the 
Receptaculites Limestone, a lithic unit some 270 
to 400 metres above the Spinella yassensis 
Limestone, and dated as mid-Emsian (probable 
conodont zone inversus-laticostatus). In their 
checklist of Australian fossil fish, Long and 
Turner (1984) listed other Australian occurrences 
of Ligulalepis toombsi (see Fig. la): the Broken 
River Group, Queensland, Early Devonian sites in 
New South Wales, and the Thangoo Calcarenite 
of the Canning Basin, Western Australia (a single 
scale, identified as cf. Ligulalepis, Turner et aL, 
1981). Scales of a second species, L. yunnanensis 
Wang and Dong 1989, have been reported from 
the Late Silurian of China. Wang and Dong, in 
their description of this species (1989: 203), state 



that the scale peg, socket, and ligula are absent. 
Schultze (1968) includes these three features in 
his generic diagnosis, and so the affinities of L. 
yunnanensis are open to question. 

In Schultze's description of L. toombsi, the 
species is not allocated to a family but is left in the 
'bucket' grouping, the palaeonisciforms. There is 
currently no satisfactory classification of the 
palaeonisciforms (Schultze & Bardack 1987), and 
the terms palaeonisciform, palaeoniscoid and 
palaeoniscid are often used interchangeably. Even 
within the family Palaeoniscidae, genera that are 
included by some workers have been placed in 
entirely different families and even different orders 
by other workers. The term 'palaeoniscid' can be 
particularly confusing: it is sometimes used in 
referring to members of the family Palaeoniscidae 
(e.g. Esin 1990), or of the order Palaeoniscida (e.g. 
Moy-Thomas & Miles 1971), or of the superorder 
Palaeonisci and the order Palaeoniscidae 
(Kazentseva 1964). In this paper, the term 
'palaeoniscoid' refers to fishes of the order 
Palaeoniscida exclusive of the deep-bodied 
platysomoids. As it is so difficult to categorise 
articulated specimens, it is understandable that a 
genus such as Ligulalepis, which is known only 
from isolated scales, should have been identified 
no more closely than 'palaeoniscoid 1 . 

This paper describes several new types of 
Ligulalepis scales from the Early Devonian of 
NSW. Ligulalepis scales of various forms may be 
attributed to particular regions of the body, 
following the pattern of squamation described by 



176 



C. BURROW 



la 



Q Broker* River Group 




Present Stddy 



ee Group 




c~~i 



1b 



FIGURE l.a) Map of Australia depicting known sites of Ligulalepis toombsi scales, b) Localities of sites with L. 
toombsi scales in study - Locality A: sites C091, C092, C624, C625. B: sites C600, C608. C: site C287. Position of 
sites based on Australian Topographical Map Series 1:250 000 Narromine and Nymagee. Map adapted from Pickett 
and McClatchie (1991: Fig. 1). 



TABLE 1. Distribution of scale types at different sites studied 



Sites 


Tl 


T2 


T3 


T4 


T5 


T6 


T7 


T8 


T9 


Trundle Beds 

SiteC287(7br + 
5 lepidotrichia) 


1L 
4R 


2L 


5L 
3R 


1L 


1R 


1 


1L 
1R 


1L 
2R 





Mineral Hill 
Site C625(34 br) 


5L 
5R 


5L 
2R 


3L 
3R 


2L 
1R 


1L 
2R 


2 


1L 


1L 





Mineral Hill 
Site C092(7 br) 








2R 




















Mineral Hill 
Site C624 


























1L 


Mineral Hill 
Site C628 





1R 


1L 




















Mineal Hill 
SiteC091 





1L 























Jerula Formation 
Site C600- only 
2 broken scales 




















T=Type, L=Left, R=Right, 


br=broken. 



















SCALES OF A PALAEONISCOID 



177 



Esin (1990) in the Permian palaeoniscid 
Amblypterina costata Eichwald. 



MMMC = Fossil Collection of the Mining and 
Mineralogy Museum, Sydney. 



Material 



Scales of ligulalepis toombsi 



The study is based on 67 complete scales which 
are classifiable into 9 types, 50 pieces of scales, 
and 5 lepidotrichia. In addition, there are several 
pieces of dermal bone, bearing the chevron pattern 
of ornament characteristic of ganoid scales. Scales 
described in this paper came from residue 
limestone samples treated with acetic acid by Dr 
John Pickett (Geological Survey NSW) in 
searching for conodonts. The parent sites are in 
the Gleninga Formation of the Mineral Hill Group, 
Trundle Beds of the Trundle Group, and the Jerula 
Formation; all being in the Murda Syncline of 
western NSW, of pesavis and/or sulcatus 
conodont zones (Pickett 1992, Pickett & 
McClatchie 1991; see Fig. lb). Table 1 details the 
distribution of scale types for the various localities; 



Schultze (1968) attributed four scale forms to 
Ligulalepis toombsi. Of these, the scale selected 
as holotype differed from all known palaeoniscoid 
scales in having its rostro-dorsal corner developed 
into a prominent tongue-shaped projection 
(described as 'loffelformig', or spoon-shaped, by 
Schultze 1968: 346; see Figs 2,3). In nominating 
a scale as a holotype, it is conventional and 
appropriate to choose an example from 'the 
anterior and middle parts of the lateral surface of 
the fish body on which the majority of 
morphological features are distinctly manifested' 
(Esin 1990: 93). In the case of palaeoniscoids, 
scales from other areas of the body may be referred 
to the same species by virtue of qualitative 
features - the form of the anterior margin of the 



r dorsal edge 



Crown View 



posterior margin of 
bony base 



Basal View 



secondary keel 




0.2mm 



FIGURE 2. Ligulalepis toombsi Schultze cf. Holotype scale from area A, with descriptive nomenclature as used in 
text - a, crown view, b, basal view, c, rostral edge-on view. 



178 



C. BURROW 




FIGURE 3. L. toombsi scales from: The mid-flank region of area A/B: a, crown view; b, basal view (ref. no. 
MMMC01926). Area B: c, crown view; d, basal view; e, conjectured basal view of interlocking adjacent scales (ref. 
no. MMMC01927). Area A, from region immediately behind the shoulder girdle and on the lateral line (note indent 
on anterior margin of scale): f, crown view; g, basal view (ref. no. MMMC01928). Area B/C: h, crown view; i, basal 
view (ref. no. MMMC01929). Area F: j, crown view; k, basal view; I, basal view of interlocking adjacent scales (ref. 
no. MMMC01930). 



SCALES OF A PALAEONISCOID 



179 



free field, the denticulation of the caudal margin 
of the scale, and detail of the sculptured ornament. 
These features were described for L. toombsi by 
Schultze (1968), as follows: anterior margin of the 
free field has an ornament of small knobs, with 
main ornament cover extending behind them; 
posterior margin serrated; numerous pores 
scattered over the ganoine surface; free field 
exhibits the chevron pattern typical of ganoine 
scales. 



This study reveals considerable variation in the 
shape of the rostro-dorsal process, from tongue- 
shaped to pennant-shaped. Consequently, the term 
'spoon-shaped' may be too narrow a description; 
in this paper it is described merely as the 'rostro- 
dorsal process'. About 22% of scales possess this 
process. Scales of Ligulalepis yunnanensis, as 
illustrated by Wang and Dong (1989), appear to 
lack this process, though this is possibly an effect 
of breakage (G. Young, pers. comm.). 




FIGURE 4. L. toombsi scales from: Area E: a, crown view; b, basal view (ref. no. MMMC01931). Area G/H: c, 
crown view; d, basal view; e, basal view of interlocking adjacent scales (ref. no. MMMC01932). Central ridge line - 
fulcral scale: f, crown view; g, lateral view; h, basal view; i, crown view of adjacent scales (ref. no. MMMC01937). 
AreaD: j, crown view; k, basal view (ref. no. MMMC01933). Area B/F: note dorso-caudal/ventro-rostral orientation 
- 1, crown view; m, basal view (ref. no. MMMC01939). Lepidotrichial scale: n, crown view; o, basal view (ref. no. 
MMMC01934). 



80 



C. BURROW 




SCALES OF A PALAEONISCOID 



181 



Schultze (1968) stated that the holotype scale is 
from the anterior half of the body. The height to 
length ratio of the scales is maximal for each row 
at the centre of the flank. By analogy with scales 
on articulated specimens of other Palaeozoic 
palaeoniscoids (e.g. Gross 1953, Gardiner 1984, 
Stamberg 1989, Burgin 1990, Esin 1990), nearly 
all overlapping scales in this study would have 
been oriented on the fish's body in rows directed 
from rostro-dorsal to caudo- ventral, except for 
scales from the latero- ventral angle. 



Descriptions 

Nine types of scales are attributed to L. toombsi, 
together with lepidotrichia. Descriptive 
terminology (see Fig. 3) follows Schultze (1968) 
and Esin (1990). 

Type 1 (Figs 3a-e, 5a) 

Scales are roughly rhomboidal, with a straight 
anterior margin except for the rostro-dorsal 
process. The dorsal peg is high with a pointed 
apex, matched by a ventral deep triangular socket. 
Primary and secondary keels are both well 
developed (as in Fig. 3). The ornamented free- 
field extends about two-thirds the length of the 
scale, and overhangs the caudal margin of the 
scale's bony base. The depth:length ratio varies 
between approximately 3/1 and 3/2 (excluding peg 
and flange). Rarely, these scales may have pores 
for the lateral line canal. Scale ornament resembles 
that described in the holotype. 

Type 2 (Figs 3f, g) 

Scales generally similar in shape to those of 
type 1, but lacking a rostro-dorsal process. The 
dorsal peg and ventral socket are very weakly 
developed. Primary and secondary keels are 
prominent. The length of the unornamented 
depressed field is about one-third to one-quarter of 
scale length on the rostral margin, and from one- 
eighth to one-quarter the depth of the scale, on the 
dorsal margin. The depth:length ratio varies 
between approximately 3/1 and 2/1. Scale 
ornament resembles that of the holotype, though 
lacking the triangular "knobs" along the rostral 
margin of the free field. 



Type 3 (Figs. 3h,i) 

Scales of roughly rhomboidal form, with a 
straight anterior margin, and a small rostro-dorsal 
process. The dorsal peg and socket are weakly 
developed, whereas the primary and secondary 
keels are pronounced. The length of the 
unornamented depressed field is negligible, on 
both the anterior and dorsal margins. The 
depth:length ratio is about 2/1, and scale ornament 
resembles that of type 2. 

Type 4 (Figs 3j — 1, 5b) 

Scales of skewed, roughly lozenge-shaped, 
outline, with an attenuated and sharply pointed 
rostro-dorsal corner. The peg and socket are 
present and very broad-based. The keels are 
parallel to the anterior margin of the scale, which 
is oriented obliquely (rostro-dorsal to caudo- 
ventral) relative to the horizontal ornament of the 
scale. The depressed field is about one-quarter the 
length of the scale, and the depth:length ratio is 
about 1/2. Ornament resembles that of the 
holotype, including the "knobs" along the rostral 
margin of the free field. 

Type 5 (Figs 4a,b, 5c) 

Scales of rhomboidal shape, without peg and 
socket. There are no keels, but the base of the 
scale has a dark central swelling. The depressed 
field on the dorsal and rostral margins is about 
one-quarter to one-fifth the length of the scale. 
The scales are slightly deeper than long, and their 
ornament is typical, aside from (like types 2 and 
3) lacking the "knobs" along the rostral margin of 
the free field. 

Type 6 (Figs 4c-e, 5d) 

Almond-shaped to diamond-shaped scales. Peg, 
socket and keels are all lacking, but the base of 
the scale has a dark central swelling. The 
depressed field extends around the dorsal, rostral 
and ventral margins. The depth:length ratio is 
about 2/5. The ornamented surface extends to a 
point overhanging the caudal edge of the base. 
Scales lack ornament 'knobs' along the margins 
of the free field. 



Type 7(Figs4f-i, 5e) 

Scales of sub-rhomboida 



form with a 



FIGURE 5. Ligulalepis toombsi scales - (scale bar = 0.1mm) - a, Broken type 1, with large lateral line pore opening 
(ref. no. MMMC01938); b, Crown view, type 4 (ref. no. MMMC01930); c, Crown view, type 5 (ref. no. 
MMMC01935); d, Basal view, type 6 (ref. no. MMMC01932); e, Crown view, type 7 (ref. no. MMMC01937); f, 
Crown view, type 8 (ref. no. MMMC01934); g, 2 lepidotrichia: basal view (left) and lateral/crown view (right) (ref. 
no. MMMC01936); h, Fragment of dermal bone, showing chevron pattern. 



182 



C. BURROW 



transversely arched base. Peg, socket and keels 
are absent, but once again the base has a slight 
dark swelling located centrally. The depressed 
field extends along the dorsal, rostral and ventral 
margins. Ornament resembles that in scales of 
type 6, but raised at approximately 60° from the 
base and pointing dorso-caudally. 

Type 8 (Figs 4j,k, 5f) 

Narrow scales of lenticular outline. There are no 
pegs, sockets or keels, and the base is of uniform 
thickness. The depressed field extends from the 
rostral point dorsally to the caudal point, but is 
very narrow. The depth: length ratio varies from 
about 1/3 to 1/6. The ornamented surface is almost 
smooth, aside from a few shallow striations, and 
does not form an overhang. 



Lepidotrichia (Figs 4n,o) 

These have a long, narrow 'bread-loaf shape, 
with a longitudinal furrow in the middle of each 
side. There is a central peg and socket on some 
lepidotrichia, of the full thickness of the scale, and 
a groove running between these on the base of the 
scale. The lepidotrichia range in length from about 
0.6mm to 1.0mm, and their width is about 0.1mm. 
The flat upper surface is relatively smooth, though 
pores are present and the surface is lightly striated. 
In contrast to the body scales, the ornament may 
extend onto the peg. The 'unnamed palaeoniscoid 
scales' illustrated by Giffin (1980, Fig. 11) appear 
to be examples of lepidotrichia from L. toombsi. 



Discussion 



Type 9 (Figs 41, m) 

Scales of 'bent' rhomboidal form, without peg 
or socket. There is a primary keel and a weak 
secondary keel. The depressed field is about a 
third the length of the scale. The depth:length ratio 
is about 2/1, and the long axis is oriented in a 
dorso-caudal to rostro-ventral direction. The 
ornament is typical, though it is noticeably more 
worn on the ventral half; it extends beyond the 
caudal margin of the scale base. 



Scale variation according to body area 

Having categorised the scales into these 
different types, the question now is - from which 
sections of the body might they have come? 
Morphology of scales is known to vary in 
consistent fashion from area to area of the body in 
articulated specimens of palaeoniscoids (Gross 
1953, Gardiner 1984, Long 1988, Esin 1990). 




FIGURE 6. Diagrammatic outline of fusiform palaeoniscoid fish, showing distribution of major areas with 
morphologically distinct scales, a, area behind pectoral girdle; b, front half of flank; c, rear half of flank; d, tail 
(anterior to fin); e, in front of dorsal fin; f, ventral; g, surrounding base of dorsal fin; h, surrounding base of anal fin I 
& II, dorsal unpaired scale rows. 



SCALES OF A PALAEONISCOID 



183 



Consequently nine types of scales here described 
for L. toombsi may be assigned to specific areas of 
the body. The following designations are modelled 
on the scheme of variation described by Esin 
(1990) for Amblypterina costata Eichwald (see 
Fig. 6). 

There is a decrease in the depth of scales from 
area A to area D, and from the median line of the 
fish to its dorsal and ventral margins. Thus the 
scales with greatest depth/length ratio should be 
assigned to area A. However, the first row of 
scales behind the shoulder girdle would not 
include forms depicted in Fig. 3, with their 
rostrally directed process, and it is for this reason 
Schultze (1968) assigned scales of type 2 to the 
area immediately behind the shoulder girdle. The 
scales would be expected to be well articulated, 
being in a relatively inflexible area of the body; 
scales of type 2 fit the criteria for scales along the 
rostral margin of area A, overlapping scales of 
type 1. 

Scales of type 1 probably came from areas A 
and B, overlapping type 3 scales towards the B/C 
margin. 

Scales in area C would not be expected to be 
highly articulated with each other, as interlocking 
devices (keels, pegs, sockets and, in the case of L 
toombsi, rostro-dorsal processes) are weakly 
developed or absent on scales from more flexible 
areas of the body (Esin 1990). Scales of type 3 
best fit the criteria exhibited by scales of this 
moderately flexible area in articulated 
palaeoniscoids. 

Scales in the flexible area D lack articulations 
and are non-imbricating. Scales of type 8 would 
seem to fit here, becoming more elongated 
towards the tail fin. 

Scales from area E are expected to have a broad 
low peg, with keels being weakly developed or 
absent, and a wide depressed field (Esin 1990). 
Scales of type 5 match these criteria best. 
Although they lack a peg, they are of a shape 
intermediate between that of scales of type 2 and 
type 7. 

With regard to area F, the length of the wholly 
ventral scales of Palaeozoic palaeoniscoids 
described to date is invariably greater than their 
height, and they have poorly developed pegs and 
sockets, with an extended rostro-dorsal corner and 
a wide rostral depressed field, for secure 
anchoring in the skin. Scales of type 4 come from 
this area. Schultze (1968: Plate I, figs 6a,b) stated 
that such scales were definitely from the ventral 
region. Scales of type 9, with a bent shape and 
worn ornament on the lower half must surely have 



come from the flank/ventral angle of the body. 
These were the only scales to have a rostro-ventral 
to dorso-caudal orientation, indicating a ventro- 
lateral scale row directional inversion. 

Scales at the base of the fins are expected to be 
small relative to neighbouring scales, and to show 
bilateral symmetry (Esin 1990). Scales of type 6 
fit these criteria, and by comparison with scales 
figured in Esin (1990) they most likely came from 
the base of one of the paired fins (areas G and/or 
H). 

Scales of type 7 are probably ridge or fulcral 
scales (areas I and II of Esin 1990). Presumably 
these were aligned in staggered pairs along the 
'ridge' line. No bilaterally symmetrical scales 
have been observed. In other Devonian, and more 
recent, palaeoniscoids there is wide variation in 
the distribution of paired and unpaired fulcral 
scales. Of those species found in Australia, the 
Late Devonian Howqualepis rostridens Long 
1988 has unpaired dorsal and ventral fulcral 
scales on the posterior half of the body; Mimia 
toombsi Gardiner and Bartram 1977 has unpaired 
fulcra along the whole body dorsally, and ventrally 
from the pelvic fin to the tail; Moythomasia 
duagaringa Gardiner and Bartram 1977 has 
unpaired fulcra in front of all unpaired fins except 
the anal fin (Gardiner 1984). According to 
Kazantseva (1976) fulcra were initially paired in 
palaeoniscoids; i.e., in the primitive condition 
there were no symmetrical fulcra. As Ligulalepis 
is one of the oldest palaeoniscoids found in the 
fossil record (Late Silurian of China - Wang & 
Dong 1989), it would be expected to lack 
symmetrical (i.e. unpaired) fulcral/keel scales. 

Although lepidotrichia are rare, they appear to 
be of a distinctive form. Like the body scales, their 
structure indicates they were more tightly locked 
together than those of other fishes. 



Functional interpretations 

Scales with high depth:length ratios, from the 
mid-flank region, are quite strongly curved (Fig. 
2c). Perhaps this is of significance in relation to 
the extra interlocking device - i.e., the rostro- 
dorsal process. The 'abnormal' palaeoniscoid 
Cheirolepis (see Pearson & Westoll 1979, Pearson 
1982) has micromeric squamation and also lacks 
the typical interlocking devices (pegs and sockets) 
of its contemporaries, the stegotrachelid 
palaeoniscoids. Ligulalepis scales of the same 
form as the holotype have the highest depth to 
length ratio reported for any Palaeozoic 



84 



C BURROW 



palaeoniscoids, and their strong curvature implies 
a low scale number per scale row, perhaps 10 or 
fewer. Scales of platysomoids have a high 
depth:length ratio, but other features of L toomhsi 
scales would preclude them from belonging to a 
platysomoid (e.g. their strong curvature, and their 
rostro-dorsal processes). A box-shaped or circular 
transverse section would be ruled out by the fact 
that mid-flank scales show the greatest curvature. 
A fusiform shape is indicated by elimination of 
these other possible shapes, and by the correlation 
shown in this paper between the scale types 
observed for L. toomhsi with those observed for 
more recent fusiform fishes. 

There is apparently a correlation between the 
size of scales and the degree of their interlinking. 
For example, the Cretaceous palaeoniscid 
Cteniolepidotrichia probably comprises two 
species (Poplin & Su 1992), one having deep 
scales equipped with pegs and sockets, and the 
other having smaller squarish scales without pegs 
or sockets. The body covering of large scales 
indicates that Ligulalepis did not swim in the 
shark-like fashion envisaged for Cheirolepis by 
Pearson and Westoll (1979). Peg and socket 
articulations would have constrained dorso-ventral 
flexibility, while the anterior processes limited the 
lateral flexibility of the body. The large interlocked 
scales probably indicate that swimming involved 
low amplitude undulations. The scale rows would 



have acted to brace the sides of the trunk, 
preventing twisting of the body. Contemporaneous 
fish with non-micromeric squamation included 
many placoderms, whose early forms had armour 
extending onto the trunk so that the front half of 
the body was stiffened. Gottfried (1991) suggested 
that this stiffening, for placoderms and deep- 
scaled fish, also assisted air ventilation by recoil 
aspiration, as among polypterids. In any case, 
Ligulalepis toomhsi appears to have had the least 
flexible trunk region of any known palaeoniscoids. 
It is not possible to infer the likely habitat of L. 
toomhsi by analysis of the other microvertebrate 
remains in the same samples. At some sites, 
acanthodian and placoderm scales were most 
abundant, while at other sites thelodont scales 
predominate. This diversity of faunal associations 
may indicate that L. toomhsi had an extensive 
ecological range, from near shore environments to 
further out on the continental shelf. 



Acknowledgments 

I wish to thank Dr John Pickett and the Geological 
Survey of NSW for providing the samples used in this 
study. Dr Sue Turner assisted with techniques for sorting 
the samples and gave helpful advice; Mr Rick Webb 
helped with SEM photography, Dr Gavin Young 
(AGSO) provided information, and Dr. Tony Thulborn 
gave helpful advice and criticism. 



References 



BURGIN, T. 1990. Palaeonisciden (Osteich- 
thyes:Actinopterygii) aus dem Unteren Rotliegenden 
(Autunien) der Nordschweiz. Eclogae Geologicae 
Helvetiae 83(3): 813-827. 

CAMPBELL, K. S. W. & BARWICK, R. E. 1988. 
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phylogenetic hypotheses. Geological Magazine 
125(3): 207-227. 

ESIN, D. N. 1990. The scale cover of Amblypterina 
costata (Eichwald) and the Palaeoniscid taxonomy 
based on isolated scales. Paleontological Journal 2: 
90-98. 

GARDINER B. G. 1984. Relationships of the 
palaeoniscoid fishes, a review based on new 
specimens of Mimia and Moythomasia from the 
Upper Devonian of Western Australia. Bulletin of the 
British Museum (Natural History), Geology series 
37: 173-428. 

GARDINER, B. G. & BARTRAM, A. W. H. 1977. The 
homologies of ventral cranial fissures in 
osteichthyans. Pp. 227-245 in 'Problems in 
Vertebrate Evolution'. Eds. S. M. Andrews, R. S. 
Miles & A. D. Walker. Academic Press: London. 



GIFFIN, E. M. 1980. Devonian Vertebrates from 
Australia. Postilla 180: 1-15. 

GOTTFRIED, M. D. 1991. A new deep-scaled 
'Palaeoniscoid' from the Upper Pennsylvanian of New 
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32A. 

GROSS, W. 1953. Devonische Palaeonisciden-Reste im 

Mittel- und Osteuropa. Paldontologische Zeitschrift 
27:85-112. 

KAZANTSEVA, A. A. 1964. Subclass Actinopterygii. 
Pp. 410-775 in 'Fundamentals of Paleontology. 
Volume XI - Agnatha and Pisces'. Ed. D. V. 
Obruchev. (Translated from Russian by Israel 
Program for Scientific Translations, 1967, Jerusalem). 

1976. Fulcra and keel scales in 

palaeoniscids. Paleontological Journal 10: I 13-115. 

LONG, J. A. 1988. New palaeoniscoid fishes from the 
Late Devonian and Early Carboniferous of Victoria. 
Memoirs of the Association of Australasian 
Palaeontologists 7: 1-64. 

LONG, J. A. & TURNER, S. 1984. A Checklist and 
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in 'Vertebrate Zoogeography and Evolution in 



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Australasia 1 . Eds. M. Archer & G. Clayton. Hesperian 
Press: Carlisle Western Australia. 1203pp. 

MOY-THOMAS, J. A. & MILES, R. S. 1971. 
'Palaeozoic Fishes'. 259 pp. Chapman & Hall: 
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PEARSON, D. M. 1982. Primitive bony fishes, with 
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PEARSON, D. M. & WESTOLL, T. S. 1979. The 

Devonian actinopterygian Cheirolepis Agassiz. 
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337-399. 

PICKETT, J. W. 1992. Review of selected Silurian and 
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(GS 1992/024) 

PICKETT, J. W. & McCLATCHIE, L. 1991. Age and 
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POPLIN, C. & SU, D. 1992. Cteniolepidotrichia 
turfanensis n.g. n.sp., a bizarre Palaeoniscoid (Pisces, 
Acinopterygii) from the Cretaceous of Xinjiang, 
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Geologie und Palaontologie, Monatsheft 8: 469-486. 



SCHULTZE, H-P. 1968. Palaeoniscoidea-schuppen aus 
dem Unterdevon Australiens und Kanadas und aus 
dem Mitteldevon Spitzbergens. Bulletin of the British 
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SCHULTZE, H-P. & BARDACK, D. 1987. Diversity 
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(Actinopterygii, Pisces) from the Pennsylvanian 
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STAMBERG, S. 1989. Scales and their utilisation for 
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(3): 255-269. 

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Edwin Sherbon Hills'. Ed. R. W. Le Maitre. 
Distributed by Blackwells, Carlton. 

TURNER, S., JONES, P. J. & DRAPER, J. J. 1981. 
Early Devonian thelodonts (Agnatha) from the Toko 
Syncline, western Queensland, and a review of other 

Australian discoveries. Bureau of Mineral Resources 
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WANG, N. Z. & DONG, Z. Z. 1989. Discovery of late 
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192-206. 



CAPE HILLSBOROUGH : AN EOCENE - OLIGOCENE VERTEBRATE 
FOSSIL SITE FROM NORTHEASTERN QUEENSLAND 

GregMcNamara 



Summary 

The Hillsborough Basin is a narrow graben that parallels the modern coast between Prosperine and 
Mackay, north Queensland. At Cape Hillsborough, just north of Mackay, Hillsborough Basin 
deposits outcrop. Here a succession of volcanic and volcaniclastic rocks, oil shales and ostracodite 
named the Cape Hillsborough Beds are well exposed. The Cape Hillsborough Beds are informally 
subdivided into a dominantly volcanic succession, termed the Cape Hillsborough volcanics, 
separated by an angular unconformity from an underlying ostracodite and oil shale succession 
termed the Wedge Island beds. 



CAPE HILLSBOROUGH: AN EOCENE - OLIGOCENE VERTEBRATE FOSSIL SITE 

FROM NORTHEASTERN QUEENSLAND 

GREG MCNAMARA 



MCNAMARA, G. 1994. Cape Hillsborough: an Eocene - Oligocene vertebrate fossil site from 
northeastern Queensland. Rec. S. Aust, Mus. 27(2): 187-196. 

The Hillsborough Basin is a narrow graben that parallels the modern coast between 
Prosperine and Mackay, north Queensland. At Cape Hillsborough, just north of Mackay, 
Hillsborough Basin deposits outcrop. Here a succession of volcanic and volcaniclastic rocks, oil 
shales and ostracodite named the Cape Hillsborough Beds are well exposed. The Cape 
Hillsborough Beds are informally subdivided into a dominantly volcanic succession, termed the 
Cape Hillsborough volcanics, separated by an angular unconformity from an underlying 
ostracodite and oil shale succession termed the Wedge Island beds. 

The Wedge Island beds contain vertebrate fossils including ubiquitous teleost bones, turtle 
(chelid) bones and crocodile scutes. 

Palynoflora extracted from Wedge Island beds oil shale indicates a Middle Eocene age based 
on correlation with the Nothofagidites asperus Zone. A mean K/Ar age of 32.5 ± 0.4 Ma on the 
Cape Hillsborough volcanics provides a minimum Early Oligocene age for the underlying 
sediments and the Cape Hillsborough fossil fauna. The fauna is therefore derived from one of 
the better dated Early Tertiary vertebrate fossil sites in Australia. 

G. McNamara, Department of Earth Sciences, James Cook University of North Queensland, 
Townsville 481 1. Manuscript received 26 August 1993 



Cape Hillsborough is a coastal promontory on 
the northeastern Queensland coast located 
between Mackay and Proserpine (Fig. 1). 
Hillsborough Basin sediments and volcanics are 
well exposed here. The Hillsborough Basin is a 
narrow graben structure within Palaeozoic 
volcanic, plutonic and sedimentary rocks. It trends 
south-southeast from Proserpine under Repulse 
Bay and the Hillsborough Channel to near 
Mackay. 

The Cape Hillsborough area was probably a 
horst relative to other parts of the Hillsborough 
Basin graben which resulted in a thinner 
stratigraphic pile developing and a sequence of 
sediments and volcanics which are not readily 
correlated with sequences elsewhere in the basin 
(Slessar 1970). 



Geology 

The Cape Hillsborough Beds 

At Cape Hillsborough, outcrop is dominated by 
about 300m of felsic and mafic pyroclastics and 
lava flows overlying oil shale and limestones 
collectively termed the Cape Hillsborough Beds. 
The type locality for the Cape Hillsborough Beds 



(Clarke et al. 1968) takes in all the north - 
northeastern cliff exposures from Cape 
Hillsborough to Andrews Point within the Cape 
Hillsborough National Park (Fig. 2). 

The Cape Hillsborough area encompasses all 
known exposures of Cape Hillsborough Beds 
(Slessar 1970). They consist mostly of rhyolitic 
and basaltic lavas and pyroclastics but comprise a 
complex interfingering of volcanic and 
volcaniclastic rocks deposited during a short lived 
episode of pyroclastic volcanism. Small exposures 
of underlying sediments occur at low tide 
revealing approximately 15 m of succession. 
Slessar (1970) interpreted these sediments as the 
upper part of a more extensive oil shale sequence 
with which the volcanics are unconformable. 

Remapping has confirmed the volcanics and 
sediments are not conformable but are divided into 
two units by an angular unconformity. The 
sediments below the unconformity are informally 
termed the Wedge Island beds after the key 
outcrop at Wedge Island. They encompass other 
sequences recorded from Donna Bay, Mackay Oil 
Prospecting Syndicate (MOPS) 4 and MOPS 5 
drill holes, GSQ Proserpine 1-2RA drill hole and 
other outcrops mapped by Slessar (1970) but not 
relocated by the author (Fig. 2). The overlying 
volcanics are informally termed the Cape 



G. MCNAMARA 




FIGURE 1 . Regional geology and location of Cape Hillsborough. 



Hillsborough volcanics and include all Tertiary 
volcanics, volcaniclastics and epiclastics 
outcropping stratigraphically above the Wedge 
Island beds in the Cape Hillsborough area (mostly 
within Cape Hillsborough National Park; Fig. 2). 

Slessar (1970) noted the sediments contained 
fossil plant material, invertebrates and fish bone. 
Other vertebrate fossils from the Wedge Island 
beds are described here. Tuffaceous sediments 
within the Cape Hillsborough volcanics have 
yielded angiosperm leaves and other plant material 
(Clarke et al. 1968). The existence of fossiliferous 
tuffs and volcaniclastics in the sequence raises the 
interesting possibility that vertebrate fossils may 
also be present within the volcanic pile. The author 



located a boulder-sized piece of silicified wood 
[JCU FI2517] in a pyroclastic breccia halfway 
between Cape Hillsborough and the GSQ 
Proserpine 1-2RA drill hole (Fig. 2). This 
indicates significant quiescent periods occurred 
between eruptive events that would have allowed 
for the return of vertebrates to the area over the life 
of the volcanic activity. 



Stratigraphic relationships and age of the 
Wedge Island beds 

Slessar (1970) correlated the outcropping 
Wedge Island beds with the oil shales and 



CAPE HILLSBOROUGH FOSSIL SITE 



189 



Donna Ba 



Cape Hillsborough 




Wedge Island 

Causeway 



§ Basal sediments mapped by 
Slessar but not relocated 



a Basal sediments 



Palaeozoic basement 



Cape Hillsborough Beds 



FIGURE 2. Geology of Cape Hillsborough National Park (after Slessar 1970). 



sandstones intersected in the wells MOPS 4 and 
MOPS 5 drilled in 1956. The wells were drilled in 
the southern part of the Cape Hillsborough study 
area. The stratigraphic relationship between the 
sediments and volcanics is not delineated by these 
drill holes because they only intersected the 
sediments (Clarke et al. 1968). However, there is 
no doubt Slessar' s correlation is correct. The 
location of MOPS 4 and MOPS 5 is less than 50 
m from an (unrelocated) outcrop of the sediments 
(Fig. 2). Bedding attitudes at this locality and 
Donna Bay indicate that the sequence dips under 
the volcanics. There is no evidence of a faulted 
contact. Intersections recorded for MOPS 4 & 
MOPS 5 do not include bimodal volcanic units 
between the sedimentary sequence and the 
Palaeozoic basement, further confirming the view 
that the sediments are basal to the volcanics rather 
than the volcanics being faulted in from below the 
sediments. 

In 1971 a stratigraphic bore (GSQ Proserpine 
1-2RA) was put down by the Queensland Mines 



Department at Cape Hillsborough (Fig. 2). The 
Wedge Island beds were intersected approximately 
30 m below the volcanics, continuing to a depth of 
453 m, thus confirming the interpretation of 
Slessar. The sediments in this section were logged 
as interbedded mudstone, shale, siltstone, 
sandstone and oil shale (Swarbrick 1974). 

There is only one outcrop, located at the 
northern end of the Causeway (Fig. 2), where the 
contact between the sediments and volcanics is 
exposed. Slessar (1970) interpreted the contact at 
the Causeway as an angular unconformity but 
remapping of the site in 1987 showed it to be a 
fault. Dip measurements indicate that the Wedge 
Island beds are folded with the north-dipping limb 
of a gently plunging anticline truncated against 
subhorizontal basalt. The exposed fault is steeply 
dipping and trends almost parallel with the strike 
of the north-dipping beds (Fig. 3). 

Both rock types are brecciated at the contact but 
no sense of movement is apparent. Assuming the 
folded sediments are expressing a tectonic event 



190 



G. MCNAMARA 



Wedge Island 




100m 
Andrews Point 



Cape Hillsborough volcanlcs 



Wedge Island beds 



FIGURE 3. The geology of Cape Hillsborough between Andrews Point and Wedge Island and the location of the 
main fossil deposit at the Causeway. 



CAPE HILLSBOROUGH FOSSIL SITE 



191 



that pre-dates the volcanism, it follows that the 
fault block containing them has been uplifted 
relative to the volcanics. The causeway outcrop 
sits at a marginally higher elevation than other 
outcrops of sub-volcanic sediments. It seems 
unlikely that all other sub-volcanic outcrops would 
be faulted-in to near the same level. A more likely 
explanation is that the other outcrops express 
undisturbed relationships and the causeway 
sequence, although repositioned by faulting, has 
not experienced large vertical displacement. 

Slessar did not find contacts for any of the other 
outcrops of Wedge Island beds (Fig. 2) but 
assumed they were contiguous on the basis of 
coincident dips and the interpreted angular 
unconformity at Wedge Island. However, the 
stratigraphic relationship between the Wedge 
Island beds and the Cape Hillsborough volcanics 
is not in question, only the size of the hiatus 
between them. The timing of tectonism with 
respect to the volcanism and the stratigraphy 
revealed in GSQ Proserpine 1-2RA clearly 
indicate that the Wedge Island beds, and hence 
the vertebrate fossils they contain, are sub- 
volcanic. The folding of the Wedge Island beds at 
the Causeway indicates an angular unconformity 
does exist between the sediments and the Cape 
Hillsborough volcanics even though the faulted 
contact does not. 

Hodgson (1968) analysed samples from MOPS 
4 and MOPS 5 for spores and pollen. MOPS 4 
produced a good yield of well preserved pollen 
whereas MOPS 5 failed to produce a microflora. 
The palynoflora from MOPS 4 included 
Haloragacidites harrisii, Nothofagus cf N. 
deminuta and lnaperturopollenites sp. and thus 
Hodgson (1968) concluded that the sample was 
probably Lower Tertiary in age. Hekel analysed 
samples collected by V. Palmieri at Donna Bay 
and material from MOPS 5. For Donna Bay 
samples, he recorded elements of the 
Cupanieidites orthoteichus zonule, including its 
nominate species, indicating a Paleocene to Early 
Eocene age and probably constrains the maximum 
age as Late Paleocene (Hekel 1972; Foster 1980). 
Slessar (1970) listed a flora provided by Hekel for 
MOPS 5 and noted it contains elements of the 
Cupanieidites orthoteichus zonule together with 
elements of the Myrtaceidites eugeniiodes and 
Gambieriana edwardsii zonules then believed to 
be of Paleocene age in South Australia (Harris 
1971). Hekel (1972) concluded that the Cape 
Hillsborough Beds must be Paleocene to Middle 
Oligocene in age by correlating them with the 
Hillsborough Basin sediments known at 



Proserpine from extensive drilling (eg AEQ 
Proserpine 1 (Hutton 1980)). 

Samples were collected at the Causeway by the 
author [JCU 36309 - 36320] in an attempt to 
improve palynological data. The resultant 
palynofloras were assessed by consultant 
palynologists McEwan-Mason and Wagstaff, 
Melbourne and also Neville Alley, South 
Australian Department of Mines and Energy. The 
yield of palynomorphs was poor but their 
preservation was fair to good (N. Alley pers. 
comm.). The initial assessment (McNamara 1993) 
indicated a palynoflora that provided no useful 
biostratigraphic information (N. Alley pers. 
comm.). Subsequent assessment provided a more 
useful palynoflora. 

The dominant taxa in these samples are 
Haloragacidites harrisii and Araucariacites 
australis. Malvacipollis diversus, Nothofagidites 
deminutus, N. heterus, Rhoipites spp., 
Podocarpidites spp. and Triorites cf. T. 
orbiculatus (Foster 1982) are all present in 
significant numbers. 

It is noted that Proteacidites frequency and 
species diversity is relatively low, with the most 
consistent of the seven species present being 
Proteacidites pachypolus. Proteacidites 
kopiensis, P. reticulatus, Triporopollenites 
gemmatus, Anacolosidites sectus and 
Nothofagidites falcatus are present. Rare 
occurrences of Diporites aspis, Crassoretitriletes 
vanraadshooveni, Malvacearumpollis man- 
nanensis, Polyodiaceoisporites retirugatus and 
Polypodiidites usmensis are also noted. 

The palynofloras lack the characteristics of 
Australian Paleocene and Early Eocene 
assemblages, missing the typical common to 
locally abundant taxa. The palynofloras are also 
not typical of Late Eocene and Oligocene 
assemblages (N. Alley pers. comm.). Significantly, 
they lack Triorites magnificus which is a reliable 
indicator of early Late Eocene time, its first 
appearance defining the base of the Middle 
Nothofagidites asperus Zone of the Gippsland 
Basin (Stover & Partridge 1973) which coincides 
with the base of Late Eocene. 

The first appearance of Nothofagidites falcatus, 
present in the samples, defines the base of the 
Lower Nothofagidites asperus Zone (Stover & 
Partridge 1973) which corresponds with the early 
Middle Eocene. This indicates a Middle Eocene 
age for the Wedge Island beds. A correlation with 
the N. asperus Zone is supported by the lack of 
Triorites magnificus and the presence of 
Anacolosidites sectus which is restricted to the 



92 



G. MCNAMARA 



Lower N. asperus Zone. Further support is given 
by the presence of Proteacidites kopiensis and P. 
pachypolus whose range ends in the Lower N. 
asperus Zone (Dudgeon, 1983). 

Dudgeon (1983) correlated the central 
Queensland Yaamba Basin deposits with the 
Nothofagidites asperus Zone using a very similar 
palynoflora. It too was derived from an oil shale 
succession. This suggests the Rundle, Yaamba 
and Condor oil shale successions are 
penecontemporaneous and all Eocene in age. 
However, diachronous species ranges between 
Queensland basins and the Gippsland Basin, 15° 
to the south, must be considered. Foster (1982) 
and Dudgeon (1983) concluded that apparent 
upward range extensions of species in Queensland 
did not invalidate the use of southern Australian 
data. This does not diminish the problem of 
diachronous ranges produced by latitudinal effects 
on the temporal distribution of flora. Latitudinally 
controlled zones occur earlier in northern 
sequences rather than later (N. Alley pers. 
comm.). 

Sluiter (1991) noted that floral differences 
between Eocene Lake Eyre sediments and 
southern sites are not great, even though Lake 
Eyre is 9° further north (about half way between 
the southern sites and Cape Hillsborough). Sluiter 
(1991) suggested this lack of difference may be 
due to a weak equator to pole gradient for this 
period (Kemp 1978). If Sluiter is correct it may 
mean there is little latitudinal bias in Eocene 
species distribution and the Middle Eocene age 
correlation for the Wedge Island beds is 
reasonable. 

McDougall and Slessar (1972) dated six 
samples (with concordant results) from the Cape 
Hillsborough volcanics, with a mean age of 32.5 ± 
0.4 m.y. (Early Oligocene). This age is consistent 
with the stratigraphic relationship between the 
Cape Hillsborough volcanics and the Wedge 
Island beds and the angular unconformity between 
them. 

The age of the volcanics is commonly quoted as 
the age of the whole of the Cape Hillsborough 
Beds which is misleading. A considerable hiatus, 
perhaps as much as 12Ma, separates the Wedge 
Island beds and the Cape Hillsborough volcanics. 

The new floral data from Wedge Island concurs 
with previous palynological assessments of an 
Early Tertiary age. It refines the age of the beds 
and is significant for two reasons. Firstly, it 
indicates the deposit is much older than the 
Oligocene age of the overlying Cape Hillsborough 



volcanics. Secondly, it clearly demonstrates a 
Middle Eocene age for the vertebrate fossils. 



Stratigraphy and sedimentology of the Wedge 
Island beds 

At the Causeway outcrops of Wedge Island 
beds delineate a small anticline gently plunging 
towards the northwest. The beds inclined at 20" on 
the northern limb are exposed at low tide and 
represent 15m maximum vertical thickness 
(Figure 3). At Donna Bay the thickness accessible 
above the low water mark is approximately 2m 
and the facies associations are distinctly different. 
Both sequences have abundant bone preserved 
within the coarser facies. 

Four facies are recognised: 

Facies 1 : Ostracodite. A lime-rich, cream- 
coloured, sometimes cross-bedded sandstone with 
occasional pebble sized, well rounded, 
allochthonous clasts and rip-up clasts. Sand grains 
are principally ostracod skeletons but terrigenous 
grains of quartz, feldspar and lithics are present. It 
is generally carbonate cemented and fossil bones 
(light brown) are present throughout [JCU 36315, 
36316, 36318, 36320 & 36321]. 

Facies 2 : Muddy limestone. Cream-coloured, 
massive siltstone/mudstone. Induration varies 
from fully lithified to virtually unlithified 
dependent on the degree of carbonate cementation 
[JCU 36314, 36317 & 36319]. 

Facies 3 : Grey-brown shale (oil shale). Finely 
laminated. Oily efflorescence sometimes apparent. 
Carbonaceous plant impressions present 
throughout. Poorly indurated, sea washed sections 
appear massive [JCU 36309, 36310, 36311, 
36312 & 36313]. 

Facies 4 : Brown pebbly sandstone. Variable 
grainsize (very fine-very coarse) both laterally and 
vertically. Grains dominantly terrigenous but 
ostracods very abundant. Iron oxide staining 
common. Fossil bone (jet black) present 
throughout. Carbonate cement [JCU 36322]. 

The Causeway facies association 

Two distinct facies groupings are present in the 
Causeway section. The base of the logged section 
is a monotonous sequence of Facies 3. It is at least 
7.5 m thick, extends out beyond the low tide mark, 
and crops out very poorly (Fig. 4). It contains 
traces of plant material and has yielded pollen and 
fungal spores but no bone was found. 

The oil shale is replaced up section by a 



CAPE HILLSBOROUGH FOSSIL SITE 



193 



CM 



2 




*tf- 



<N- 



oil shale 



C 

5 



(D 



C - Crocodile 
F-Fish 
P - Pollen 
T- Turtle 



."^W^ 



Fault breccia 

Faulted contact 

limestone 

C "Croc-rock" 

T F 

no outcrop 

no outcrop 
limestone F 



FIGURE 4. Logged section CHI: Wedge Island beds, 
Wedge Island. 

sequence of interbedded muddy limestone and 
ostracodite but the contact is obscured (Fig. 4). 
Many of the ostracodite outcrops have sharp bases 
and appear to grade into the muddy limestone 
facies. They may represent a fining upwards 
association but in other outcrops the distinction 
between the two facies is more pronounced and no 
fining upwards association is apparent. The 
maximum thickness expressed by this association 
is 4.5 m (Fig. 4) but is variable along strike. 

The Donna Bay facies association 

The thickness available for logging at Donna 
Bay during low tide amounts to approximately 
2m. More is exposed above the intertidal zone but 
it is mostly covered by a beach boulder bed. 
Consequently it was decided there was little value 



in logging the section. The facies association at 
this locality does not include either of the 
Causeway limestone lithologies but the rocks are 
carbonate rich. Instead there is an interbedded 
sequence of grey-brown shale and brown pebbly 
sandstone. The beds of both types grade between 
each other over a short distance but sharp contacts 
are uncommon. Carbonate is present as abundant 
ostracod valves and as cement. 

Fossil bone is found in both facies at this 
locality but principally in the brown pebbly 
sandstone. 



Wedge Island beds environment of deposition 

Oil shales develop in anoxic environments such 
as deep freshwater and hypersaline lakes 
(Demaison & Moore 1980). Green & Bateman 
(1981) concluded that the Hillsborough Basin is 
structurally similar to the East African rift system 
and that the large anoxic lakes found in that 
system are a good modern analogue for the 
depositional environment of the Hillsborough 
Basin although marine incursions must have also 
occurred. Green & Bateman (1981) propose a 
hypothetical 'Condor Lake', at a palaeolatitude of 
approximately 45° south, developed in an 
intramontane graben. Sedimentation kept pace 
with prolonged subsidence, allowing for the 
accumulation of very thick fluvio-lacustrine 
sequences of oil shales and other sediments. 
Dinoflagellates, acritarchs and alginite B in some 
sections together with anhydrite and gypsum 
crystals indicate brackish to marine conditions 
suggesting the basin was sometimes open to the 
sea (Green & Bateman 1981) whereas the alga 
Pediastrum sp. in other sections indicates a 
freshwater environment (Foster 1980). 

The near-shore environment was dominated by 
a temperate rainforest flora, similar to forests 
containing Nothofagus fusca type trees found in 
Tasmania and New Zealand today. Temperatures 
were probably mild; averaging 17-1 8°C (Sluiter 
1991). The forest was probably relatively closed 
and probably grew very close to the depositional 
basin (Martin 1978). This type of temperate forest 
is ideal habitat for a wide range of birds, 
mammals and other animals today and there is no 
reason to think that it would have been any less 
hospitable during the Eocene. 

Slessar (1970) identified ostracods from Donna 
Bay as Limnocypridae, Bisulcocypris and 
Metacypris and noted that they indicate a non- 
marine environment but provided no stratigraphic 



194 



G. MCNAMARA 



insights. Ostracods collected by the author from 
Wedge Island were too damaged and too well 
cemented to identify (P. De Deckker pers. comm.). 
Freshwater ostracods identified from the Rundle 
and Duaringa oil shale sequences, which are 
similar to the Wedge Island sequence, suggest 
deposition in a shallow (approximately 1 m deep) 
oxygenated lake (Fleming et ai 1979). 

The ostracodite is not a conventional limestone. 
It is a fluvio-lacustrine deposit where the dominant 
grain type is paired ostracod valves [JCU 36321]. 
Other, non-biogenic, non-carbonate terrestrial 
grains are common. The bulk of the carbonate in 
the rock is cement infilling the ostracod valves 
and occluding pore-space. The valves themselves 
may have provided some soluble carbonate for 
precipitation through abrasion and dissolution 
prior to deposition but they are only calcified 
chiton, not carbonate. Pebble sized terrigenous 
clasts and muddy/silty limestone rip-up clasts are 
well rounded. Cross-bedding is evident in many of 
the boulders strewn on the causeway but not so 
obvious in outcrop. Fish bones are entirely 
disarticulated while the turtle bones are carapace 
fragments and carapace-plastron sections which 
do not readily disintegrate, even in energetic 
streams. 

The ostracodite therefore, most likely represents 
the reworking of fluvial terrigenous input and 
lacustrine accumulations of ostracod, fish, turtle, 
crocodile and ?other vertebrate debris. Assuming 
a freshwater to brackish water environment 
prevailed, a wave-washed lake margin with 
numerous small streams debouching into a ?very 
shallow to shallow lake seems an appropriate 
environment to generate such a sediment. The 
ostracodite-muddy limestone facies transitions 
may even be recording transgressive-regressive 
lake levels with the anoxic oil shale facies 
representing the deeper, off shore, sections of the 
lake. Rip-up clasts may indicate occasional sub- 
aerial exposure of the marginal lake floor or 
proximity to a stream flowing into the lake. 

The ostracodite-limestone facies is not recorded 
elsewhere in the Hillsborough Basin (Green & 
Bateman 1981; Green et al 1984) but is recorded 
in the oil shale sequences of the Rundle Formation 
further to the south (Coshell 1983). The Rundle 
Formation contains sequences of oil shale and 
ostracodite that are virtually identical to the 
Wedge Island sequence. Coshell (1983) proposed 
a shallow lacustrine environment for the 
deposition of the Rundle Formation sediments and 
noted that depositional cycles included 
transgressive and regressive events causing 
reworking of lake sediments by wave action and 



brecciation through sub-aerial exposure. 
Ostracodite horizons were recognised by Coshell 
(1983) as representing very shallow lacustrine 
conditions that only rarely were associated with 
sub-aerial exposure. 

Palaeontology 

The Wedge Island local fauna 

Vertebrate fossils at Wedge Island are confined 
to the ostracodite and muddy limestone facies. 
Bone material is commonly an unidentifiable hash 
but there are abundant spines and vertebrae which 
clearly indicate that the bulk of this material is 
from fish. 

The Wedge Island local fauna is described 
below. Unfortunately none of fossils are 
sufficiently diagnostic to allow the taxon it 
represents to be identified in detail. The specimens 
have not been figured for this reason. The concept 
of the local fauna follows Tedford (1970). 
Catalogue numbers refer to specimens catalogued 
and held at James Cook University. 

Crustacea 
Ostracoda 

Ostracod shells form a significant clastic 
component in the ostracodite facies but can also 
be found in oil shales and siltstones at Donna Bay 
[JCU 36323, 36324] (Slessar 1970). 

Mollusca 
Gastropoda 

Casts of naticoid gastropods are common in the 
Ostracodite and muddy limestone facies at Wedge 
Island and are very common in some horizons at 
Donna Bay but cannot be more accurately 
identified [JCU 36325]. 

Osteichthyes 
Teleostei 

Fish spines, vertebrae and other 'fish hash' are 
ubiquitous and can be found in all of the non-oil 
shale facies. No articulated material or diagnostic 
bone has been found [JCU 36326, 36327]. 

Reptilia 
Chelidae 

Turtle carapace and plastron sections and 



CAPE HILLSBOROUGH FOSSIL SITE 



195 



fragments are the most common fossils in the 
deposit after fish. They probably represent chelid 
bone but the ornamentation and suturing features 
are not diagnostic and no further refinement in the 
identification is possible (Gaffney, pers. comm.; 
Gaffney 1991) [JCU F12509, F12510, & F1251 1]. 

Crocodylidae 

Several distinctly crocodilian dermal scutes 
were found at Wedge Island in an outcrop dubbed 
'croc-rock' - CHI: 11 m (Fig. 4). While quite 
characteristic of the family they are of no help in 
further identification (Molnar, pers. comm.) [JCU 
F12512&F12513]. 

Problematica 

Coprolites were also found in outcrop at croc- 
rock. While these particular specimens have no 
diagnostic value they are large enough to have 
come from a crocodile [JCU F12514]. One 
specimen however, is reminiscent of stools left by 
large aquatic birds such as swans [JCU F12515]. 



Provenance and bias of the Wedge Island local 
fauna 

A shallow lacustrine environment would have 
provided favourable habitats for many vertebrates 
but may not have been conducive to the 
preservation of their remains. Oil shales and other 
anoxic sediments that preserve plant material are 
not necessarily suitable for preserving bone. There 
is very little bone present in the oil shale outcrops 
at Wedge Island. The casts of fish vertebrae 
reported by Green and Bateman (1981) indicate 
bone dissolution is taking place locally within the 
Condor oil shales. 

Conversely, the chemical conditions for bone 
preservation within the limestones were probably 
quite good, especially if cementation occurred 
rapidly after deposition. However the ostracodite 
was in fact a fluvio-lacustrine deposit involving 
the transport and reworking of sedimentary 
components including bones. Such an 
environment was destructive of articulated 
remains and of individual bones through 
mechanical abrasion. 

Terrestrial animals would have had their 
remains abraded during fluvial transport to the 
lake and through wave action on beach and deltaic 
bedforms (Napawongse 1981). Aquatic animal 
remains would also have been subject to wave 
action but not the dual action of fluvial transport 



and lacustrine reworking. Therefore, despite a 
favourable habitat, the depositional environment 
of the ostracodite facies mitigated against the 
preservation of bone of terrestrial origin. 

Additionally, biogenic degradation of bone may 
be expected even before any bone transport begins. 
Teeth are likely to survive longer than many bones 
but anything that remains will be greatly diluted 
in the sediment load relative to the remains of 
aquatic taxa. 

Not surprisingly then, all the bone found to-date 
derives from aquatic species with the most 
abundant fossil taxa (fish) simply reflecting the 
natural aquatic abundance probably present in the 
depositional system and the number of individual 
spines, vertebrae and other bones to be found in 
the average teleost. Turtle carapace bones and 
crocodile scutes are dense, robust bones likely to 
survive mechanical abrasion. In addition, turtles 
and crocodiles are likely to be common taxa in a 
lacustrine environment and each individual has a 
large number of such bones or scutes. 



Recommendations for future work at cape 
hillsborough 

It is unlikely that since fish, turtle and 
crocodiles were present, the lacustrine 
environment and surrounding terrestrial habitat 
was hostile to other vertebrates. Even though not a 
single bone or tooth from a non-aquatic taxon was 
found, the importance of this site remains 
undiminished. 

Given the small number of Palaeogene 
vertebrate fossil sites and the degree of age control 
on the Wedge Island beds, one of the best-dated 
Early Tertiary vertebrate bearing units in 
Australia, this unit has good potential for 
important future discoveries. The potential for 
vertebrate fossil discoveries in the Cape 
Hillsborough volcanics should also be explored. It 
is recommended the succession be routinely 
prospected. 



Acknowledgments 

The Queensland National Parks Service provided 
permission for fossil collection and support on site. 
James Cook Univeristy provided Special Research Grant 
funding to allow the research to proceed. Professor Bob 
Henderson, James Cook University, is thanked for his 
comments on earlier drafts. Neville Alley, S.A. Dept. of 
Mines and Energy, is especially thanked for his thorough 
review of this work and his reworking of the 
palynological data at short notice. 



96 



G. MCNAMARA 



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vertebrate fossil deposits. Unpublished MSc thesis 
James Cook University, Townsville. 

NAPAWONGSE, P. 1981. A taphonomic study of bird 
bone degradation in fluviatile environments. 
Unpublished honours thesis, Monash University. 

SLESSAR, G. 1970. The geology of Cape Hillsborough, 
Mackay region. Unpublished B.Sc. honours thesis, 
James Cook University, Townsville. 

SLUITER, I. J. 1991. Early Tertiary vegetation and 
climates, Lake Eyre region, northeastern South 
Australia. Pp. 99-118 in: The Cainozoic in Australia: 
A re-appraisal of the evidence. Ed. by M. Williams, 
P. De Deckker & A. P. Kershaw. Geological Society 
of Australia Special Publication No. 18. 

STOVER, L. E. & PARTRIDGE, A. D. 1973. Tertiary 
and Late Cretaceous spores and pollen from the 
Gippsland Basin, southeastern Australia. Special 
Publications of the Geological Society of Australia 
4: 55-72. 

SWARBRICK, C. 1974. Oil shale resources of 
Queensland. Geological Survey of Queensland 
Report No. 83, Brisbane. 

TEDFORD, R. 1970. Principles and practices of 
mammalian geochronology in North America. 
Proceedings of the American Palaeontological 
Convention 1969: 666-703. 



THE MIOCENE OSCILLATION IN SOUTHERN AUSTRALIA 



Brian McGowran & QianyuLi 

Summary 

Warm times are moister times and times of higher sea level. They are represented by more 
extensive stratigraphic records, richer fossil assemblages, and more precision and confidence in 
correlation and age determination. The Miocene profile in southern Australia is a profile of 
episodically advancing seas from the late Oligocene to the early middle Miocene, then a 
pronounced retreat into the late Miocene. In parallel is the chronological distribution of three warm 
periods - in the Janjukian stage, the mid-Longfordian stage (early Miocene), and then the twin- 
peaked Miocene climatic optimum of the Batesfordian-Balcombian stages of the earliest middle- 
Miocene, 16-15 million years ago. After that, there is a pronounced cooling. That rise and fall at 10 
years scale is the Miocene oscillation. The planktonic foraminiferal record at lakes Entrance in East 

■7 

Gippsland supports the concept of a rise and fall in sealevel and climate at 10 scale with the 
superimposition of an oscillation at higher frequency - 10 years. The match with the global 
scenario is good. As well, the most pronounced changes in the fossil record are seen within the 
Miocene climatic optimum, not at the subsequent chilling and fall in sea level. The terrestrial biotas 
are expected to reveal in due course this three-part Miocene succession : (i) episodically rising 
trends, Janjukian to late Lonfordian ; (ii) Miocene climaxes, Batesfordian and Balcombian ; (iii) 
drying and cooling, Bairnsdalian to Mitchellian. The Australian regional stages have a role in the 
geochronology of Cainozoic biogeohistory ; their boundaries should accord with major natural 
breaks in the stratigraphic record. 



THE MIOCENE OSCILLATION IN SOUTHERN AUSTRALIA 



BRIAN MCGOWRAN & QIANYU LI 



MCGOWRAN, B. & LI, Q. 1994. The Miocene oscillation in southern Australia. Rec. S. Aust. 
Mas. 27(2): 197-212. 

Warm times are moister times and times of higher sea level. They are represented by more 
extensive stratigraphic records, richer fossil assemblages, and more precision and confidence in 
correlation and age determination. The Miocene marine profile in southern Australia is a profile 
of episodically advancing seas from the late Oligocene to the early middle Miocene, then a 
pronounced retreat into the late Miocene. In parallel is the chronological distribution of three 
warm periods — in the Janjukian stage, the mid-Longfordian stage (early Miocene), and then the 
twin-peaked Miocene climatic optimum of the Batesfordian-Balcombian stages of the earliest 
middle Miocene, 16-15 million years ago. After that, there is a pronounced cooling. That rise 
and fall at 10 7 years scale is the Miocene oscillation. The planktomc foraminiferal record at 
Lakes Entrance in east Gippsland supports the concept of a rise and fall in sealevel and climate 
at 10 7 years scale with the superimposition of an oscillation at higher frequency— 10 fi years. The 
match with the global scenario is good. As well, the most pronounced changes in the fossil 
record are seen within the Miocene climatic optimum, not at the subsequent chilling and fall in 
sea level. The terrestrial biotas are expected to reveal in due course this three-part Miocene 
succession: (i) episodically rising trends, Janjukian to late Longfordian; (ii) Miocene climaxes, 
Batesfordian and Balcombian; (iii) drying and cooling, Bairnsdalian to Mitchellian. The 
Australian regional stages have a role in the geochronology of Cainozoic biogeohistory; their 
boundaries should accord with major natural breaks in the stratigraphic record. 

Brian McGowran and Qianyu Li, Department of Geology and Geophysics, The University of 
Adelaide, South Australia 5005. Manuscript received 2 June 1993. 



We have undertaken a research program based 
on the Neogene foraminiferal succession in 
southern Australia. As the continent 'drifted' 
north, subsequent to rapid Australia/Antarctica 
separation commencing in the middle Eocene, it 
moved into lower latitudes whilst global climate 
was deteriorating. Its margin was washed by a 
series of transgressions. In sharp contrast to the 
trailing passive margin in the south, the northern 
margin began, during the Oligocene, to accrete 
terrains — the latest events in a history of tectonic 
activity stretching back to the Palaeozoic. 

There are essentially three reasons for 
undertaking this research. One is the relevance of 
a well-developed, mid-latitude, neritic record of 
calcareous strata to oceanic drilling to the south of 
Australia in the Ocean Drilling Program — for the 
neritic-oceanic interaction bears heavily on the 
problems of reconstructing a global sea level curve 
and of clarifying links between palae- 
oceanographic change and palaeoclimatic change. 
A second stimulus is that the rich neritic record of 
foraminifera, ostracodes, bryozoa and echinoids, 
in addition to the molluscs which have long 
attracted attention, is a splendid resource for 
studying macroevolution, meaning organic 
evolution at time scales of 10* and 10 7 years. Yet a 



third reason is that the terrestrial biotas of ancient 
Australia have left records susceptible to 
prolonged argument on the two basic and critical 
questions: What is the 'age' of the assemblage? 
What was the 'environment 1 of its community? By 
supplying the links between continental landform 
and biotic evolution on the one hand and global 
change and its geochronological framework on the 
other, the marine strata at the continental margins 
can provide something of a reference for 
addressing both questions. It is this third reason 
that preoccupies us here, although it is impossible 
to isolate it from the others. 



Consilience of induction: the prime strategy of 
stratigraphy 

The weakest point in virtually all philosophers' 
systems is the place of history in their scenarios of 
science and its doing. Among their biological 
achievements, three luminaries can count some 
incisive discussion of the fact that not all science 
is circumscribed by the laws and immanent 
properties of the physical sciences: Simpson 
(1963, 1970), Mayr (1982), Gould (1986, 1989). 

We can use an example from correlation at 



198 



B. MCGOWRAN & Q.LI 



terrestrial plants 
temperature °C 



5 18 OtoPDB 



meters to PSL 

o o o g g 

o o o ^ £ 

co c\] i- o T . 




20 
l_ 



30 



40 
I 



latitude °N 
neritic molluscs 



50 



O 

o 



I 

o 

o 



deep water temperature 
-* falling 



global 
sea 
level 

falling 



FIGURE 1. Correlation of time series, adapted from McGowran (1991). Left, terrestrial plant assemblages from the 
western U.S.A. yield temperature estimates (Dorf 1955) and molluscan faunas give estimates of latitudinal shift in 
isotherms (Durham 1950). Centre, a composite 8 ls O signal from benthic foraminifera in the deep ocean (Shackleton 
1985); the thermometer becomes distorted from the Miocene onward by the ice effect. Right, the putative curve of 
global sea level (Haq et al. 1987). Note the close general matches among the various kinds of reconstruction: sea 
level falls as climate deteriorates through the Cainozoic Era. There are mismatches: the molluscan study missed the 
Miocene reversal; the major fall in Oligocene sea level lagged behind the spectacular fall in isotopic temperature by 
the duration of the early Oligocene. Also included are the named ingressions and transgressions of southern 
Australia— benchmarks in the marine record of the southern continental margin. 



geological time scale of 10 fi years to make that 
point. Consider the elegant scenario of Cainozoic 
change, usually described as global climatic 
deterioration, in Figure 1 (McGowran 1991). We 
have long known that the Eocene was a time of 
global warmth: uniformitarian evidence for that 
generalization consists of the occurrence of 
crocodiles, palms, molluscs, corals at higher 
latitudes than are attained by their modern 
counterparts (e.g. Lyell 1867). Using the same 
kind of strategy — extrapolation back from modern 
biogeographic faunal distribution, including what 
was known of the environmental requirements of 
coastal molluscs — Durham (1950) showed that 
marine isotherms for the western margin of North 
America have been retreating equatorwards since 
the Eocene. Similarly, Dorf (1955) could employ 
terrestrial floras as 'thermometers of the ages' to 



give a generalized curve for 40-50N latitude in 
the western U.S.A. Both strategies are strong, but 
both are open to quibbling about the reliability of 
fossil leaves or of shells as indicators of past 
climatic states. We add Shackleton's (1985) 
generalized oceanic bottomwater curve of d 18 
values from benthic foraminifera. That sort of data 
too could be and was criticized as susceptible to 
diagenetic alteration, vital effects, salinity changes 
in the reservoir — criticisms in the same 
uniformitarian vein as for the fossil curves, and to 
be taken seriously. And yet the three curves have a 
powerful mutual similarity! If quite disparate data 
from the terrestrial, neritic and pelagic realms of 
the biosphere show such a good mutual match 
through geological time, then the chances that we 
are seeing real climatic changes are suddenly 
much better than they were for each of the data 



MIOCENE OSCILLATION 



99 



sets in isolation. The mutual reinforcement of 
shells, leaves and isotopes is greater than the sum 
of the parts. Adding a curve of putative global sea 
level (Haq et al. 1987) to the array, we see a 
strengthening by further persuasive correlations. 
Sea level has fallen as temperature has fallen, from 
the same high point in the early Eocene. (There is 
a nagging mismatch where the major cooling 
leads the biggest fall in sea level by the duration 
of the early Oligocene — also heuristic because it 
demands explanation.) 

That lesson in mutual reinforcement by 
chronological correlation is at Cainozoic time 
scales. We can amplify the scenario for the 
Miocene to make the same point by anticipating a 
major theme of this paper (Fig. 6). McGowran 
and Li (1993) have correlated two kinds of curve 
independently to a revised chronology (W. A. 
Berggren, pers. comm., 1992). One is the oceanic 
isotopic evidence for the Mi glaciations at 10 6 
years' scale, rising then falling on the trend at 10 7 
years' scale (Wright et al. 1992); and the other is 
the Exxon sea level curve, also at both 10 7 and 10 ft 
years' scale (Haq et al. 1987). The match at both 
scales is remarkable, even though the independent 
correlation to the integrated chronology leaves 
several mismatches at 10 6 years' scale: coolings 
should fit sequence boundaries, not maximum 
flooding surfaces, either because of glacioeustatic 
lowering of sea level, or because lower sea levels 
enhance continental-type climates. Thus the 
mismatches can be targeted for scrutiny and a 
fine-tuning of the correlations. 

In both of these examples it is the chronological 
correlation of disparate data that counts for most. 
This strategy has a noble ancestry and yet has 
been curiously unacknowledged (Gould 1986, 
1989). Charles Darwin, 'so keenly aware of both 
the strengths and limits of history, argued that 
iterated pattern, based on types of evidence so 
numerous and so diverse that no other 
coordinating interpretation could stand — even 
though any item, taken separately, could not 
provide conclusive proof — must be the criterion 
for evolutionary inference' (Gould 1986). As 
Gould notes, the 19thC philosopher of science 
William Whewell called this strategy of 
coordinating different lines of evidence to form a 
historical pattern, 'consilience of induction'. 
(No matter that Whewell — coiner of 
'uniformitarianism' and 'catastrophism' — was an 
essentialist philosopher (Mayr 1982) with little 
grasp of Darwin's iconoclastically historical 
science; or that he not only rejected the thesis of 
'On the origin of species...' but banned the book 
itself (Gould 1989).) 



'Consilience of induction' captures the spirit 
and the strategy of biostratigraphy. How do the 
strata and their recorded events fit together as a 
pattern in space and time, and what does that 
pattern tell us of biogeohistory? This is not an 
orderly one-way street from correlation to history, 
but rather a turbulent two-way, hermeneutic 
thoroughfare. The mutually reinforcing patterns in 
Figure 1 show the 'Miocene oscillation' or 
temporary reversal of the grand Cainozoic trend 
(McGowran, in Frakes et al. 1987) clearly, except 
for Durham's curves. And that is the simplest 
reason to reject a view of Australian Cainozoic 
history entrenched in sedimentology, 
palaeobotany, geomorphology, neritic 
palaeontology — that we have a somehow special 
climatic trajectory brought about by Australia's 
northwards drift to warmer latitudes. Rejected 
already (McGowran 1979a; in Frakes et al. 1987), 
that view has been supported again by Zinsmeister 
(1982) and Darragh (1985). The Australian 
scenario fits the global pattern too well to need ad 
hoc buttressing by auxiliary hypotheses. (Which is 
not of course to question the importance of 
Australia's present position to the modern 
climate.) 



The Oligo-Miocene 'sequence 1 

Consider Figure 2, drawn in 1977. It is a 
cartoon, a bold conjecture, of course, which 
attempts to summarize patterns of strata in space 
and time, to hammer the message that there is a 
strong parallelism in time through the full range 
of sedimentary and tectonic environments. As a 
first-order generalization the pattern still holds, 
broadly, although it emphasizes the latest Eocene 
too much and the latest-middle to earliest-late 
Eocene too little (McGowran 1989a), and the 
middle Miocene instead of the early Miocene (see 
below). Those and other corrections are less 
important than the remarkable gaps in the early 
middle Eocene, early Oligocene, and late Miocene, 
giving four Cainozoic 'sequences'. (There is some 
fossiliferous sediment of early Oligocene and late 
Miocene age; these are relative statements.) The 
gaps consist either of hiatuses well constrained by 
bounding dates, or of markedly regressive facies 
not well dated, but assuredly occupying only a 
small part of the time between good dates, or 
simply an absence or marked rarity of good 
biostratigraphic records. The theme is repeated 
and extended but thoroughly substantiated in the 
palaeogeographic and facies maps of Australia 



200 



B.MCGOWRAN&Q. LI 



Md 



SOUTHERN PASSIVE MARGIN 
offshore onshore 





ou. 










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10- 










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marginal to 
nonmarine detrital 



CENTRAL 

PLATFORM 



l — ' — r— ' — r^ — r - — 



NORTHERN ACTIVE MARGIN 

PLATFORM NEW GUINEA MOBILE BELT 

-.11— n— _n— v— • n— 11— ii— H 
...11.. v — - — - — v— V 



V 



,■■ ■ ,' i '4^^i^L+4+i++ 



"i r 



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rn 



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m <- 



PAPUAN 

ULTRAMAFIC 

BELT 



v- 



V — 



neritic 
carbonate 



I carbonate, deep water 
— other deen water — |— plutonism \/ uolcanism fTl metamorphism 



Jul. a m 5 77 



FIGURE 2. Cartoon of sediments versus time across the Australian continent, from McGowran (1979a), intended to 
show that decades of correlation and age determination — and some interpolation — yielded a pattern of 'sequences' 
bounded by hiatuses. Obviously, much lithological variety and much continent have been omitted. With some 
adjustment, the broad contrast between a richer early-middle and a poorer middle-late Miocene stratigraphic and 
fossil record still holds. 



(BMR Palaeogeographic Group 1990). Thus their 
map 'Cainozoic 2', 52-37 Ma (actually 
representing most accurately the 4 Ma time slice 
across the middle/late Eocene boundary, 42- 
38Ma) displays a generous array of marine facies 
especially on the southern margin, and nonmarine 
facies in palaeodrainages and major catchments. It 
is remarkably similar to 'Cainozoic 4', 30-10 Ma 
(actually representing most accurately the time 
across the early/middle Miocene boundary, 20- 
15Ma), and to 'Cainozoic 6', 5-2 Ma (Pliocene). 
In stark contrast are 'Cainozoic 3', 37-30 Ma 
(early Oligocene) and 'Cainozoic 5', 10-5 Ma 
(late Miocene). At those times all facies, marine 
and nonmarine, at the margins and across the 
continent, shrink almost to vanishing point. The 
cartoon in Figure 2 still holds validly as another 
way of depicting the Miocene 'sequence'. The 
sequence is the regional stratigraphic 
manifestation and signal-bearer of the Miocene 
oscillation. 



Stratigraphic correlation and classification 

Australia is antipodeal to the region where 
Cainozoic stratigraphy developed; it is separated 
from that region by the tropics; and its terrestrial 
biotas and much of the neritic biotas carry the 
stamp of prolonged isolation from the rest of the 
world. Its northern, heavily vegetated margin in 
New Guinea excepted, the continent has been 
stable, lacking the thick, rapidly accumulating 
sedimentary successions subsequently uplifted and 
deeply exposed by erosion, as in the Americas and 
the Himalaya, for example. Instead, we have 
thinner and more scattered sections with less 
outcrop and a carapace of deep weathering and 
duricrusting. Biogeography and the stratal record 
accordingly give us a double problem: to piece 
together a highly composite, egional, stratigraphic 
and biostratigraphic succession; and to correlate 
that record with the global geochronological 
standards. As if there were not complaints enough 



MIOCENE OSCILLATION 



20 



already, there is a further problem special to the 
Cainozoic at large, as outlined by Martin 
Glaessner in a masterful study of half a century 
ago: 

Biostratigraphic correlation of Mesozoic marine 
deposits is based on zones which are either 
worldwide or at least useable within the wide limits 
of a palaeo- zoo geographic province. Correlation of 
Tertiary deposits is a much more difficult problem on 
account of climatic differentiation, topographic 
isolation, and close stratigraphic subdivision of 
deposits representing a comparatively short time 
interval. No worldwide scale of fossil-zones based on 
well-defined ranges of a set of index-species exists. A 
sequence of Tertiary faunal assemblages was long 
ago established in Europe and it is not surprising to 
find that workers in other continents first turned to 
this sequence for guidance by means of direct 
comparison and correlation. As long as no scale of 
zones is available, the next higher unit in 
stratigraphic classification, the stage, must be the 
basic unit for measuring geologic time. The 
recognition of the European stages in the East Indies 
proved so difficult that a number of workers gave up 
and even condemned attempts at inter-continental 
correlations (Glaessner 1943: 52). 

In southern Australia the development of 
correlation and age determination can be divided 
into two major periods: 

(i) Until the 1940s. The biostratigraphy of the 
marine record was dominated by molluscs, as it 
was elsewhere (e.g. Singleton 1941; Darragh 
1985). Application of the Lyellian method of 
percentages of extant taxa had little success, also 
as elsewhere. Local stages were erected, partly in 
reaction to the ensuing confusion, partly to 
facilitate progress within the continent while 
correlations further afield remained contentious or 
impossible. But the stages were biological in their 
essence, based as they were on distinctive 
macrofaunas based on type localities. 'Correlation 
was by comparison of gross suites of described 
molluscs' (Darragh 1985). Until Singleton (1941) 
redefined the stages on lithological criteria, 
bringing some order into chaos, the stages were 
losing their value for two reasons — by becoming 
'nothing but convenient labels for certain well- 
known collecting grounds', and by sometimes 
being impossibly inclusive, sometimes too 
restricted to be useful (Glaessner 1951). Re- 
reading Singleton half a century later, one is 
struck by how little biostratigraphy he includes — 
fossils there are aplenty, but they are hardly central 
to the ordering and arranging of the stratigraphic 
record. And one is startled to be reminded that 
Singleton lists not one confident identification of 



pre-Oligocene strata across the continent, except 
for the limestones in the northwest where Irene 
Crespin's studies of larger foraminifera — not 
molluscs — allowed correlation with the Indo- 
Pacific letter classification erected in the 
Netherlands East Indies. 

(ii) 1940s to present. Foraminifera were described 

from the Neogene of southern Australia as long 

ago as the 1880s (Howchin, 1889) and the same 

author is acknowledged as one of the very first 

workers anywhere to describe them from a 

borehole (Howchin, 1891). Even so, the first 

substantial shift away from the molluscs and 

towards the more extensive use of foraminifera 

and lithostratigraphy in the definition and 

characterization of stages was delayed for decades, 

until Crespin (1943). But the most noteworthy 

event in retrospect was the crisp presentation of 

three zones and their significance by Glaessner 

(1951). The Hantkenina alabamensis, Victoriella 

plecte, and Austrotrillina howchini Zones were 

non-contiguous — upper Eocene, upper Oligocene, 

lower Miocene respectively — which upset some 

purists, but they surely succeeded in fulfilling 

Glaessner' s aim of providing a framework for the 

future clarifying of the problems seething in the 

morass of correlation and nomenclature in the 

southern Australian Tertiaries. As Tertiary 

correlations and age determinations progressed, 

most rapidly where microplankton could be used, 

they did so without the benefit of local stages (e.g. 

Glaessner and Wade 1958; Wade 1964; Ludbrook 

1971), or with the stages merely appended to the 

biostratigraphy (e.g. Carter, 1964; Ludbrook and 

Lindsay 1969). Indeed, Singleton (1968) became 

sufficiently sanguine about progress in clarifying 

foraminifera] and macrofaunal biostratigraphy to 

abandon the local stages altogether. McGowran, 

Lindsay and Harris (1971), attempting the first 

comprehensive correlation of the local stages to a 

modern 'global' geochronology, were not at all 

sure that we were as advanced as all that, and felt 

that we still needed a local chronostratigraphic 

focus for the diverse biostratigraphic evidence 

coming from continental to open marine 

environments. Even so, one of the authors 

(Lindsay 1981, 1985) subsequently used the 

stages extensively whereas another (McGowran 

1978-1988) did not. By and large, local stages 

have not prospered as essential 

chronostratigraphic tools in this region. In the 

most comprehensive modern surveys of the 

classical Tertiary marine fossils, the molluscs, the 

local stages are used but they do not loom very 

large (Ludbrook 1973, Darragh 1985). 



202 



B. MCGOWRAN & Q. LT 



(sub)tropical 



southern Australia 



Ma 



1 - 

2 - 
3- 

4 

5 

6 

7 

8 

9 
lO- 
ll - 
12 
13 
14 
15 
16 
17 - 
18- 
19 - 
20- 
21 - 
22- 
23- 
24- 
25 



Berggren et al., in press 



Heath & McGowran, 
1984 



Lakes Entrance succession 



epoch 



regional stages 



Pleist. 



5 



N22 



N21 
N20 



N19 
N18 



N17 



N16 



rm 



N11 



N10 



N8 



N7 



A 



N6 



N5 



N4 



P22 



-*- G. truncatulinoides 



G. margaritae 
G. tumida 

G. fenguaensis 



N. acostaensis 
P. mayeri 
G. nepenthes 
G. fonsi robusta 

G. tohsi robusta 
G. tohsi lobata 

G. tohsi tohsi 

G. peripheroacuta 
O. suturaiis 

P. glomerosa 
P, sicana 
G. birnageae 
C. dissimiiis 



G. insueta 



-r G. kugieri 

- 1 - G. dehiscens 
- 1 - G. kugieri 



G. truncatulinoides 

G. puncticulata 
G. plesiotumida 

G. conomiozea 



-*- N. acostaensis 
■>*- G. nepenthes 

-i- G. peripheroronda 



G. mayeri 
O. suturaiis 



G. sicanus 



■*■ G. trilobus 

-*- G. zealandica 

-i- G. kugieri 

J- G. woodi 



G. dehiscens 
G. semivera 



Gr. conomiozea 

Gq. dehiscens 
N. acostaensis 
G. nepenthes 

Gr. peripheroronda 



P. mayeri (acme) 
O. suturaiis 



Gr. praescituia 
Gs. trilobus 



constant G. connecta 

G. connecta 

G. woodi 

G. euapertura/ ■*■ 
constant Gq. dehiscens 
Gq. dehiscens 



P. glomerosa 

P. sicana t C. chipolensis E 

Gr. praescituia 






Pleist. 



WERRIKOOIAN 
YATALAN 

KALIMNAN 
CHELTENHAMIAN 



MITCHELLIAN 



BAIRNSDALIAN 



BALCOMBIAN 



BATESFORDIAN^ . 



LONGFOROIAN 



JANJUKIAN 



FIGURE 3. The southern Australian planktonic foraminiferal-biostratigraphic succession, shown as first and last 
appearances (McGowran & Li 1993), is correlated with the standard N-zones of the Neogene (W. A. Berggren, pers. 
comm. 1992). The regional stages are added at right. 



Figure 3 displays a correlation of the local 
stages with a succession of biostratigraphic events 
taken from the planktonic foraminifera, which in 
turn are the main basis for correlation with a 
standard. Note that local planktonic foraminiferal 
zones have been abandoned because no consensus 
was achieved in this terrain of scattered localities 
and highly composite succession within a greater 
Tasman region, necessitating continual definition 
and redefinition of the zones (McGowran, 1978, 
1986b; Lindsay, 1981, 1985). Instead, we have 
sought to compile and improve a detailed 
sequential ordering of reliable first and last 
appearances — bioevents or datums, as in Figure 3. 
At present we are in the uneasy situation of 
lacking a firm commitment either to routinely used 
(chronostratigraphic) stages or to stable 
(biostratigraphic) zones, unlike New Zealand 



which has both, with the stages well entrenched 
(Hornibrook et al. 1989). Meanwhile, one of us 
has developed the habit of thinking of the 
stratigraphic record in terms of marine 
transgressions (McGowran 1988), and this might 
be a basis for a renewed consideration of our local 
stages, as we discuss below. The more prominent 
transgressions are listed in Figure 1 . 



Warm peaks and the miocene optimum 

It is necessary to develop all possible precision 
in generalizing about the trajectory of 'climate' 
through the Miocene oscillation. The marine 
neritic record at mid-latitudes ought to reflect 
fluctuations by the comings and goings of 
pantropical elements, particularly the larger, 



MIOCENE OSCILLATION 



203 



phytosymbiont-bearing, benthic foraminifera. And 
so it has proved. A summary by McGowran (1979, 
1986a) concluded that their highly episodic or 
sporadic distribution in southern Australia was not 
merely reflecting a sporadic stratigraphic record, a 
sporadic distribution of suitable facies, or the 
vagaries of tectonic docking — the Noah's Ark 
effect. At any rate there were two well-marked 
intervals of warming with another not so clear and 
a fourth predicted to fall in the mid-Longfordian 
but missing, and now to be documented: 
(i) The Operculina-Amphistegina interval, upper 
Janjukian-lower Longfordian, latest Oligocene- 
earliest Miocene, equivalent to zones upper P22- 



N4. Actually these taxa are mostly successional 
with the interface at the first appearance of the 
planktonic Globoquadrina dehiscens and at the 
Janjukian/Longfordian boundary, as Lindsay 
(1981) pointed out in discussing the lower 
Amphistegina acme, but they do overlap, and the 
'interval' includes a window in which the 
Globorotalia kugleri group penetrates to the mid- 
latitudes. This warm pulse was not seen in the 
first of the modern Cainozoic oceanic oxygen 
isotopic profiles (Shackleton and Kennett 1975), 
which was correlated a little optimistically to the 
neritic record (McGowran 1979a). 
(ii) The addition to the synthesis: Early Miocene, 



Mannum 



Pliocene 



early 

middle 

Miocene 



early 
Miocene 



early 

Miocene 

to 

late 

Oligocene 



Loxbn 
Sands 



Morgan 
Limestone 



Finniss 
Clay 



E5S 



3333 



upper 

Mannum 

Limestone 



lower 

Mannum 

Limestone 



X* 



CO. 



ffl 



3 



331 



33 



s 



331 



33 



T-T-T 



3-3 



313 



3 



331 



13 



Xi 



33 



3=1 



3=1 



33 



iJ^ 



10 20 30 40 50% 
' ' i i j 



;9/ 0l 



180 



170 



1 60 



160 



140 



1 30 



120 ■ 








Lakes Entrance 



15% 10 



^flfa 



'*>& 



-L«fc 



vol 



3* 



3?s 



Q^connecta 



constant 
& connecta 



200 
ft 

■ 250 
- 300 

■ 350 
400 
450 
500 

V 550 
GOO 
650 
700 
750 
800 
350 

I- 900 

950 

1000 

1050 

-1100 

-1150 

-1200 



33 



33 



3 



GE 



33 



m 



33 



333 



3 



3 



3 



3 



3 



GE 



333 



3 



33 



333 



333 



33 



33 



33 



333 



333333 



533 



3=3 



33 



JrUrX 



N17 



N16 



N15 
N14 



N13 



N11 



N10 



M 



-L2 |X 



N7 



N6 



N5 



N4b 



XVI 



XV 
XIV 



XII 



XI 



VII 



IV 



N4a 
P22 



FIGURE 4. Counts of the larger benthic foraminifera Operculina and Amphistegina in sections at Mannum, western 
Murray Basin, and Lakes Entrance, Gippsland Basin. Note different scales. Planktonic foraminiferal assemblages I- 
XVI, from McGowran and Li (1993). This larger foraminiferal horizon in the upper Mannum is between the older 
Janjukian horizon (but not found in these sections in the lower Mannum), and the younger Batesfordian horizon (the 
Batesfordian is represented here by the Morgan Limestone, but the larger foraminiferal signature of the climatic 
optimum has been eroded here). 



204 



B. MCGOWRAN & Q.LI 



mid-Longfordian, upper zone N5-N6 equivalent. 
A well marked peak in the oceanic d l8 curve was 
not matched by a larger foraminiferal spike in 
southern Australia (McGowran 1979a), even 
though the oceanic peak was correlated 
satisfyingly with the Miogypsina-Heterostegina 
horizon and the horizon of giant molluscs in the 
late Eggenburgian of central Paratethys 
(McGowran 1979b). One miscorrelation of that 
time, since corrected, was to place the top of the 
Indo-Pacific stage Te 5 at planktonic foraminiferal 
zone N8 instead of N6. That error distracted 
attention from the good record of the spike on the 
western margin (Chaproniere 1975) if not the 
southern. Also, Ludbrook's (1961) record of 
Austrotrillina and Marginopora in the upper 
member of the Mannum Formation (western 
Murray Basin) was overlooked. We have corrected 
this omission with new records from the 
Gippsland and western Murray Basins (Fig. 4) 
showing spikes in the abundance of Operculum 
and Amphistegina which both seem to be coeval 
and to correlate with zone N6. It is noteworthy 
that the facies are contrasting, for that contrast 
enhances the importance of the signal as a climatic 
indicator: at Mannum the plankton counts are very 
low and Austrotrillina and Marginopora have 
been recorded whereas the plankton count is 
higher and the two genera are missing at Lakes 
Entrance. This event in southern Australia is a 
much-subdued version of the warming seen in the 
later Otaian stage in New Zealand, where tropical- 
type molluscan and larger foraminiferal 
assemblages appear in the North Island 
(Hornibrook 1990). 

(iii) Earliest middle Miocene, Batesfordian- 
Balcombian, Zones N8b-N9. This narrow interval 
contains two immigrations of Lepidocyclina, with 
Cycloclypeus in the east and Flosculinella in the 
west in the younger (Fig. 4), suggesting 
provincialism — the Austral-IndoPacific and the 
Southeast Australian Provinces (see discussion in 
Darragh 1985). We have no ready explanation for 
this provincialism although the large foraminifera 
are distinguishable ecologically and at least 
partially distinguishable in their biofacies, as 
Chaproniere (1975) showed for this region. It is 
the time of the Batesford-Morgan and the 
Balcombian marine transgressions and together 
they display easily the richest warm- water biotic 
record across southern Australia at the time of 
maximum Neogene extent of the sea across the 
continental margins (McGowran 1979a; 
subsequent discoveries in Lindsay 1981; Benbow 
and Lindsay 1988). It is the Miocene optimum at 



the climax of the Miocene oscillation. 

The Miocene optimum has been thoroughly 
confirmed in Japan and north to Kamchatka, on 
diverse marine and nonmarine biotic criteria 
(Tsuchi 1992; Itoigawa and Yamanoi 1990; 
Gladenkov 1990). The 'tropical marine 
paleotemperature spike at about 16Ma' (Itoigawa 
and Yamanoi, 1990) centres on Zone N8b but the 
concentration of data for warming to high northern 
latitudes is correlated with zones N8-N9. We see 
it in the early Badenian transgression and 
warming in central Paratethys (Steininger and 
Rogl 1983; McGowran 1979b). Emphasized here 
as the climatic optimum of the Miocene and the 
Neogene, it is labelled as climatic optimum 1 in 
the Pacific oceanic record (Barron and Baldauf, 
1990). The second major peak in the Shackleton- 
Kennett d ls O curve (Campbell Plateau) falls at the 
lower level of this bimodal optimum (McGowran 
1979a). The peak is seen at just the 'right' level in 
New Zealand in the latest Altonian and Clifdenian 
stages (Hornibrook 1978). A more recent review 
by Hornibrook (1990) repeats the observations that 
it is in Altonian time that larger foraminifera 
{Lepidocyclina, Heterostegina, Cycloclypeus, 
Planorbulinella zelandica, Amphistegina) reach 
southernmost New Zealand; that stenothermal 
warmwater Indo-Pacific mollusca in the South 
Island reached maximum diversity (Beu, 1990); 
and that these records are particularly favoured by 
the widespread shallow-neritic facies, implying 
maximum transgression. The Altonian stage 
extends by correlation from upper zone N6 to 
about the zone N8a/N8b boundary (Hornibrook et 
al. 1989) but it is clear from Hornibrook' s review 
that the 'thermal maximum' is in the upper 
Altonian and Clifdenian, i.e. correlated with zones 
N8-N9. 

All of these correlations confirm a distinct 
peaking of the Miocene oscillation, in a time 
interval in toto of one million years or even less, 
in the earliest middle Miocene. 
t(iv) Late middle Miocene, upper Bairnsdalian, 
Zone N14 equivalents. Not seen in larger 
foraminifera in southern Australia, this horizon in 
the Capricorn Basin and New Zealand was 
correlated with a Lepidocyclina horizon in Japan 
and perhaps with the early Sarmatian marine 
transgression of central Paratethys (McGowran 
1979a,b). Hornibrook (1990) suggests that this 
temporary warming followed by the extinction of 
foraminifera in New Zealand is a possibly 
significant inter-regional event. At Lakes Entrance 
we find the horizon as an acme in the planktonic 
Globigerinoides sacculifer in Assemblage XIII 



MIOCENE OSCILLATION 



205 



(see below; McGowran and Li 1993).] 
The warm horizons are shown in Figure 6. 



The section at Lakes Entrance in east 
Gippsland 

The Lakes Entrance oil shaft (Crespln 1947) 
was the one of the very first sections to contribute 
to our modern understanding of the southern 
temperate planktonic foraminiferal succession 
(Jenkins 1960). In reexamining the section, we 
have used a range of quantified criteria to 
distinguish sixteen faunal assemblages spanning 
some 20 Ma (McGowran and Li 1993). The 
criteria (Fig. 5) include changes in the relative 
abundances of five species groups approximating 
clades, the planktonic/benthic ratio, and species 
diversity. The ratio of the dominant species in the 
two dominant groups, (Globigerina woodi + 
connecta) -f (Globigerina bulloides+falconensis), 
is a good index of warmer/oligotrophic to cooler/ 
eutrophic changes (shorthand: woodi/bulloides 
ratio). An interesting plot is the actual comings 
and goings of planktonic species, sample by 
sample, thus emphasizing the presence (or 
absence) of a species, not its local range from 
lowest to highest record. 

As we emphasize elsewhere (McGowran and Li 



1993), the section falls into three parts. In 

descending order: 

(iii) Assemblages XI to XVI — collapse of the 

woodi/bulloides ratio; measures return to less 

feverish levels of oscillation. Zones upper N9-N17 

equivalents; Bairnsdalian and Mitchellian. 

(ii) Assemblages IX to X — greatest amplitudes in 

fluctuation in all measures including the woodi/ 

bulloides ratio. Zones uppermost N7 to lower N9 

equivalents; uppermost Longfordian, Batesfordian, 

Balcombian. 

(i) Assemblages I to VIII — decline in planktonic/ 

benthic; rising trends in the woodi/bulloides ratio 

peaking within the mid-Longfordian warm 

interval; stability in diversity and in incomings 

and outgoings. Zones N4 to upper N7 equivalents; 

upper Janjukian to upper Longfordian. 

There is an interesting counterintuitive 
relationship revealed in Figure 5. The positive 
spikes in the woodi/bulloides ratio probably 
indicate warmings in the upper water column, 
given what we know of the modern counterparts. 
If so, then they should match high points both in 
the planktonic/benthic ratio and in planktonic 
species diversity. It is consistently clear, however, 
that the reverse is the case. We suggest as an ad 
hoc speculation that fluctuations in wanning were 
accompanied by increases in precipitation and 
runoff from the nearby southeastern highlands, 



simple diversity . ,,, 

, , planktonics/benthics ratio 
(number of species) 

5 10 15 20 25 0.2 0.* 0.6 0.8 



cancellate-spinose/ 
spinose-non -cane el late 



outgoing species Incoming species 




FIGURE 5. Profiles of selected planktonic foraminiferal variables, based on about 15 000 specimens, Lakes Entrance 
Oil Shaft (McGowran and Li 1993). The woodi-connecta/bulloides-falconensis ratio is the dominant component of 
the cancellate-spinose/spinose-noncancellate ratio. Tie lines emphasize some of the matches, particularly between 
high woodi/bulloides, low diversity and low planktonic/benthic ratio. Hatched interval emphasizes increased 
oscillations in assemblages IX and X which span the Miocene climatic optimum. 



206 



B.MCGOWRAN&Q. LI 



causing briefly an estuarine-type, surface-water 
outflow, lowering salinity very slightly (planktonic 
foraminifera are very sensitive to salinity 
variations) and suppressing both population 
numbers and taxic diversity. Thus there may be 
here some evidence of a major increase in 
precipitation during the climatic optimum, as 
indeed one would expect. 



Correlation: global palaeoceanographic signals 

The significance of the Lakes Entrance 
planktonic foraminiferal succession becomes 
apparent in the global perspective. Figure 6 



displays correlations, independently of each other 
with the integrated Miocene time scale, of three 
factors: the Exxon sea level curve (Haq et al. 
1987); two cFO oceanic profiles which according 
to Wright et al. (1992) reveal glacial fluctuations 
labelled as the ten Mi glaciations; and 
assemblages I-XVI at Lakes Entrance. Note that 
at 10 7 years scale there are matches between sea 
level and palaeotemperature in that both rise to a 
peak at the optimum, then fall; and at 10 ft years 
scale there are some matches between glaciation 
and sequence boundary and some mismatches — a 
situation that will improve with further scrutiny. 
Note too that the Mi glaciations do not disappear 
towards the peak of the 10 7 years curve to reappear 



Ma 5 

6 

7 

8H 

9 
10-1 
11 
12 
13 

14-| 
15 
16 

17- 
18- 
19- 
20- 
21 

22- 
23- 
24" 



-C3Br 

C4 






5 18 PDB 



eustatic curves 



V 



C3 



C3A 

C3B- 



£4x 



C4A 



C4Ar 



C5 







C5r 




C5A 




C5Ar 








C5AB 


C5AC 



C5ADBH ^g 



C5B 



C5C 



C5D ™ 

C5E F 
C6 I 



C6A 


- 


C6AA 




C6AAr 




C6B 








C6C 



N17 



N16 



N15 



_N13 

N12 



N11 



N10 



f-N9- 

N8b 



N8a. 
N7 



N6 



N5 



N4b 



N4a 



P22 



XVI 



XV 



XIV 



XIII 



XII 



XI 



IX 



VIII 



VII- 
VI 



IV 




local stages 



MITCHELLIAN 
8.5 Ma 



BAIRNSDALIAN 

^COMBIAN 
_BATESFORDIAN 
17 Ma 

LONGFORDIAN 



22 Ma 

JANJUKIAN 

-24.5 Ma 



FIGURE 6. Integrated scenario for the Miocene, based on McGowran and Li (1993). Chronology at left, from W. A. 
Berggren (pers. comm., 1992). Molluscan assemblages VII-XV are from Darragh (1985). The two 8 1r O curves were 
drawn not by us but by Wright et al. (1992) by filtering a cloud of points. The long- and short-term eustatic curves 
are from Haq et al. (1987); solid lines, sequence boundaries broken lines, maximum flooding surfaces with third- 
order sequence notation. Note that those curves and the planktonic foraminiferal assemblages I-XVI from Lakes 
Entrance in Gippsland ('L.E. forams') are correlated independently with the time scale, not with each other. Fine 
tuning will improve the match between Mi glaciations and sequence boundaries (solid lines). The regional stage 
boundaries are placed as a suggestion at sequence boundaries (see text) but note that correlation of sequence 
boundaries with integrated geochronology is far from finally fixed. The three warm intervals recognized on the 
occurrence of larger foraminifera are shown hatched with the Miocene climatic optimum shown as a double, 
comprising Batesfordian and Balcombian components. 



MIOCENE OSCILLATION 



207 



on the downslope, but Mi2 is close to the summit. 
This is consistent with the identification of a 
marine glacial episode during the peak of the 
Neogene climatic optimum (=Miocene optimum) 
(Marincovich 1990). Most interesting of all is the 
sense of increased perturbation in the various 
measures in assemblages IX and X — at the 
zeniths of both climatic and sealevel trajectories 
and preceding, not accompanying, the major falls! 
It is the Miocene optimum that displays instability 
in the fossil record, not the immediately 
subsequent chilling and regression. 



Regional stages? 

Palaeontologists have long been aware that the 
Batesfordian and Balcombian stages mark major 
changes in the molluscan fossil record. That they 
also mark very brief ages, became apparent in 
their first correlation with a modern geochronology 
(McGowran et al. 1971). Although Batesfordian 
fades commonly are bryozoan limestones and 
Balcombian are marls and clays, there is more to 
it than merely fades, as Ludbrook (1973) has 
pointed out. Darragh's (1985) recognition of nine 
Miocene molluscan assemblages shows an 
interesting parallel with the planktonic 
foraminiferal record (Table 1; Fig. 6). Although 
the tabulation is incomplete, being of 'key' species 
selected for biostratigraphic significance and not 
the entire fauna, it shows a major turnover in the 
Janjukian (greatly exaggerated by a very poor pre- 
Janjukian record), followed by strong species 
incomings in both the Batesfordian and 
Balcombian with disappearances greatly 
exceeding new arrivals through the Bairnsdalian, 
then some recovery — partly facies-controlled — in 
the Mitchellian. Darragh (pers. comm. 1993) 
warns that taxonomic turnover would be subdued 



if the entire assemblages were used, even though 
his own figure (1985, Fig. 5) gives much the same 
result. The sense of major molluscan change in 
the two very short stages at the Miocene optimum 
deserves study at the full-faunal level. 

As stages, erected on molluscan assemblages in 
neritic facies, become redefined more precisely on 
planktonic/micropalaeontological criteria (which 
are external, the organisms having floated in from 
the ocean), they become rather less useful (Loutit 
et al. 1989). That is because there will always be 
a problem in recognizing precise bio-events as 
chrono-events in neritic or paralic facies, where 
diachronism or erratic preservation and recovery 
of index fossils are more likely. Instead, suggest 
Loutit et al. we should consider reorganizing 
stages as depositional sequences, for all strata 
above a sequence boundary are younger than all 
strata below it. The notion of 'warm interval' 
illustrates the problem here. The Janjukian/ 
Longfordian boundary is put at the base of 
Globoquadrina dehiscens which is also close to 
the first record of the large benthic species 
Operculina victoriensis (Carter 1964, 1990) 
which is within the warm interval and within 'the 
difficult transitional period between the 
Palaeogene and Neogene' (Ludbrook 1973). That 
necessitated prolonged discussion of the 
molluscan faunas (Ludbrook 1973) and of the 
biotic evidence for warming and its chronology 
(Lindsay 1981), and has contributed to uncertainty 
in chronology. A possible solution is 
foreshadowed in Figure 6. It would be better to 
raise the boundary to the major sequence boundary 
within the early Miocene so that the macrofaunas 
and microfaunas could be treated as a unit in local 
correlation reflecting their unity as a record of the 
'warm interval'. That shift upwards would do no 
violence to boundary stratotypes, for the stratotype 
Longfordian has no defined lower boundary in 



TABLE 1. Nine of the southern Australian molluscan assemblages, with positions in regional stage succession, and 
first and last appearances of and numbers of 'key species' (i.e. biostratigraphically significant), taken from text and 
tabulations in Darragh (1985). The assemblages are also shown on Figure 6. 



Mollusc 


assemblage 


Stage 


First 


Last 


Total 


XV 


Bunga Creek 


Cheltenhamian 


18 


10 


60 


XIV 


Rose Hill 


Mitchellian 


21 


5 


47 


xm 


Lake Bullenmerri 


Bairnsdalian 


5 


41 


67 


xn 


Gunyoung Creek 


Bairnsdalian 


4 


36 


99 


XI 


Balcombe Bay 


Balcombian 


23 


18 


116 


X 


Boornong Road 


Batesfordian 


25 


12 


105 


IX 


Fishers Point 


Longfordian 


11 


10 


90 


VIII 


Jan Juc Beach 


Longfordian 


9 


9 


90 


VLI 


Bird Rock 


Janjukian 


58 


34 


115 



208 



B. MCGOWRAN & Q. LI 



Gippsland and the Janjukian at its stratotype near 
Torquay has had its upper boundary raised well 
above the top of the Jan Juc Formation to the 
bioevent, base Gq. dehiscens. Note, however, that 
that suggestion depends entirely on searching out 
the sequence boundary 1.5/2.1 in the local strata. 

The Longfordian/Batesfordian and Batesfordian/ 
Balcombian boundaries would change little. The 
Balcombian spans an absurdly brief segment of 
the Orbulina bioseries (Carter 1964) and either it 
might be subsumed with the Batesfordian as 
substages of a Balcombian sensu lato, as Carter 
(1990) has proposed, or it could occupy cycle 2.4. 
The Bairnsdalian/Mitchellian boundary would 
change little. The point is that the concept of the 
local or regional stage would change for the better, 
and they would return to routine and widespread 
use as natural divisions of the stratigraphic record, 
should a comprehensive sequence-stratigraphic 
analysis cogently identify the boundaries. 
Meanwhile, we emphasize again that the regional 
stage boundaries shown in Figure 6 are 
suggestions, not conclusions. 



Discussion 

The Miocene succession in southern Australia 
can be compared closely with a global scenario of 
rising sealevels and temperatures from the latest 
Oligocene to the early middle Miocene, then a 
pronounced plunge in both indicators from the 
early middle Miocene to the late Miocene. The 
generalized stratigraphic patterns in Figure 2 are 
in accord with the trends in Figure 6. 
Superimposed on these broader (10 7 years) trends 
are the higher-frequency phenomena detected in 
the oceanic oxygen isotopic record, the third-order 
depositional cycles, and the local planktonic 
foraminiferal assemblages — all at comparable 
time scales in the 10 ft year band. But where we 
find numerous 'warm intervals' in the planktonic 
foraminiferal woodi/bulloides ratio at that 
frequency, we see only three warm intervals in the 
Miocene on more comprehensive criteria as well 
as at a longer time scale. They are in the 
Janjukian-early Longfordian, the mid- 
Longfordian, and the Batesfordian-Balcombian, 
the last being the twin-peaked Miocene climatic 
optimum. It is likely, but only partly indicated on 
Figure 6 and yet to be rigorously established, that 
the mini-glaciations Mil a and Milaa will fall 
between the first and second of these warm 
intervals, and Milb and Mil between the second 
and third. When it was warmer, it was wetter, and 
that too may be seen at the higher frequency in the 



woodi/bulloides ratio. 

What have these generalizations to do with the 
terrestrial biotic record? There are several 
implications: 

(i) The regional patterns have extensive good 
matches with the global patterns. Therefore, we 
would be very suspicious of arguments thai 
attempt to contrast a drier interior with a more 
humid coast, say, or a milder south with a warmer 
north. It is not that those contrasts did not exist — 
they did! — but that the chronologies and 
correlations are too weak to permit separation of 
lateral contrasts from vertical contrasts. The 
signals from the oceanic and neritic records will 
have their counterparts continent-wide in the 
various components of the environmental mosaic, 
obscure though they probably are. For example, a 
warming will be signalled coevally in three ways 
in three environments — as a d IK negative kick in 
the oceanic realm, as an immigration from the 
tropics in the (expanded) neritic realm, and as a 
change from drier to moister floras in the 
terrestrial realm. 

(ii) However, arguments about environment or 
evolution have to be at the right time scale and 
relative ages have to be correct at that scale. If 
chronological correlations of two faunas or floras 
are wrong by a million years, then comparisons 
and contrasts depending on a perceived 
contemporaneity become nonsensical, because the 
environment is oscillating at that same scale 
which is too fast for any generalization to mean 
anything. There is little to be said about (for 
example) two Mate Oligocene-early Miocene' 
faunas in this context because we simply do not 
know whether they were coeval or not. 
(Hi) However, more can be said at the longer 
scale — comparing early-middle Miocene faunas or 
floras with middle-late Miocene, for example. In 
1983-84 there were no Miocene marsupial faunas 
(other than Wynyardia) recognized as being older 
than Batesfordian in Australia (Woodburne et al. 
1985). By 1992-93, there had been a substantial 
downward revision of several, so that the early 
Miocene and late Oligocene is now populated with 
assigned terrestrial assemblages (Archer et al. 
1989, 1994; Rich et al. 1992; Megirian 1992; 
Woodburne et al. 1994). That is intrinsically a 
more likely scenario of age distributions, because 
it shifts some of the more important terrestrial 
faunas into generally warmer-wetter times, which 
are more likely to be preserved (and discovered) 
than are the biotas of cooler-drier times 
(McGowran 1986b). It is unlikely on predictable 
environmental grounds that the Riversleigh 



MIOCENE OSCILLATION 



209 



assemblages are Bairnsdalian, as Woodburne et 
al. (1985) suggested. 

(iv) There has been some disagreement over the 
broad nature of the Riversleigh environments. The 
host sediment, the Carl Creek Limestone, has 
been interpreted as a calciclastic alluvial outwash 
that could only have accumulated under relatively 
dry and perhaps even semi-arid conditions 
(Megirian 1992). That implies to Megirian that 
the Riversleigh rainforest was merely a refugium 
and the high mammalian diversities in the 
Miocene are due to the mixing-in of biotas from 
much drier distal habitats. Archer et al. (1994) 
advance several ecological and taphonomic 
arguments against this view and reiterate earlier 
conclusions that the Riversleigh assemblages 
represent biotas in low-latitude rainforests. Our 
emphasis on the distinction between phenomena 
at 10 7 and 10 fi year scales might point to a 
resolution of this conflict by accommodating both 
scenarios at the higher frequency. Thus the 
preserved faunas might be biased in their age 
distribution towards the warmer-wetter segments 
of Longfordian-Balcombian times when the 
terrain was heavily forested and the limestones 
were karsted. The sedimentological evidence for 
the depositional environment of the Carl Creek 
and equivalent limestones might be biased 
towards the Mi glacial (=cooler/drier) times of the 
same time span. Several successional and 
alternating episodes of the contrasting 
environmental regimes could easily be 
accommodated within the age range of the 
Riversleigh. 

(v) Those revisions in the correlation and age 
determination of vertebrate assemblages also 



accord better with emerging generalizations about 
Australian floras. Whereas macrofloras are more 
informative than microfloras as to environment, 
too often they are poorly constrained in their 
geological ages, so that an entire discussion can 
become diffuse as to what happened and just 
when did it happen (e.g. Hill 1992). On 
microfloral evidence mostly from the eastern 
Murray Basin, Martin (1989) concludes that the 
region largely was forested and mostly with 
rainforest prior to the middle Miocene. In the 
middle-to-later Miocene widespread rainforest 
was extensively replaced by eucalypt wet 
sclerophyll forest. The extent of moist forest and 
deep chemical weathering predictably reached 
their maximum during the Batesfordian- 
Balcombian climatic optimum. The later Miocene 
was somewhat drier, and we can now assert that a 
change toward drier sclerophyll forests would have 
begun fairly early in the middle Miocene, in the 
early Bairnsdalian, sensu Figure 6. On Martin's 
recent studies pertaining to the problem of 
Australian grasslands, Archer et al. (1994) 
comment: 'On balance, there is no evidence for 
early to mid Miocene grasslands in Australia' — a 
generalization that has the right 'feel' about it 
from our standpoint. The subsequent change in 
vegetation also entails a change towards sparser 
records, and records that are harder to date with 
confidence and precision. 



The Monterey effect? 

Why did the world warm up through the early 
Miocene, teeter on the brink for about a million 



Ma 
10-i 



15 



the Monterey effect: 
cooling by reversed greenhouse 



20 




late 
Miocene 

middle 
Miocene 



early 
Miocene 



2.0 



818Q%o 



813C%. 



FIGURE 7. The isotopic patterns demonstrating the Monterey effect operating over several million years (Vincent 
and Berger 1985). MC, MC, Monterey carbon event, initiation and termination. AA. AA ( , Antarctic ice buildup, 
initiation and termination. Eustatic curve as for Figure 6. Hatching shows the Miocene climatic optimum, identified 
on other grounds (see text and Figures 5 and 6) but most importantly fitting the pattern very well by correlation. 



210 



B. MCGOWRAN&Q. LI 



years, then plunge toward the chilly state of the 
late Neogene, never again to attain that 
greenhouse state of fifteen million years ago? The 
Monterey hypothesis of Vincent and Berger 
(1985) still fits the record well. It is a pattern 
hypothesis in which the profiles of d 13 C and d l8 
reveal cause and effect (Fig. 7). A pronounced 
positive trend in d 13 C beginning at horizon MC. 
was found in both surface and bottom oceanic 
successions, suggesting removal of light carbon 
from the oceanic reservoir (such as into the 
organic-rich Monterey Formation and its circum- 
Pacific equivalents). At a critical point, the d IS 
trend, strongly negative, went suddenly into the 
opposite direction, indicating growth of the 
Antarctic icecap and chilling of the global bottom 
waters at a CO^ drawdown, a threshold in 



reversed-greenhouse (AA). The event concluded 
after about 5m.y. at horizons MC , AA ( . It all fits 
very well chronologically with the Miocene 
climatic optimum with its attendant evidence from 
sea level and the fossil record. 



Acknowledgments 

The work was supported by an Australian Research 
Council Fellowship. We thank Michael Archer for a most 
useful preprint and, together with several other members 
of the vertebrate palaeontological community, for 
encouraging the preparation of this paper. He and Tom 
Darragh reviewed the manuscript and their comments 
were helpful. Bill Berggren generously supplied a copy 
of an unpublished Miocene time scale. 



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WORLD HERITAGE AND FOSSILS 

Mary Jinman 

Summary 

The purpose of this paper is to give a broad overview of the World Heritage Convention, to explain 
Australia's participation and to give details of the first Australian Fossil Site nomination : Murgon, 
Riversleigh and Naracoorte. 



WORLD HERITAGE AND FOSSILS 



The purpose of this paper is to give a broad 
overview of the World Heritage Convention, to 
explain Australia's participation and to give 
details of the first Australian Fossil Site 
nomination: Murgon, Riversleigh and Naracoorte. 

The Convention Concerning the Protection of 
the World's Natural and Cultural Heritage, known 
as the World Heritage Convention, was adopted 
by the UNESCO General Conference in, 1972. In 
1974, Australia became one of the first signatories. 
Since then, the number of countries that are party 
to the Convention has risen to 132. 

The World Heritage Convention aims to 
promote co-operation among nations to protect 
worldwide heritage which is of such universal 
value that its conservation is a concern of all 
people. It is intended that, unlike the seven 
wonders of the ancient world, properties will be 
conserved for all time. Member countries commit 
themselves to ensuring the identification, 
protection, conservation, presentation and 
transmission to future generations of their World 
Heritage Properties. These ideas form the very 
backbone of the Convention. 

The Convention establishes the World Heritage 
List, which identifies natural and cultural 
properties considered to be of outstanding 
universal value, and, by virtue of this quality, 
especially worth safeguarding for future 
generations. In order to qualify for inscription on 
the World Heritage List, a nominated property 
must meet specific criteria and integrity conditions 
from either a natural or cultural point of view. 

The world's natural heritage offers a priceless 
legacy. It includes sites representing major stages 
in the geological and biological history of the 
earth; outstanding examples of significant ongoing 
processes; areas of exceptional natural beauty; and 
significant natural habitats for the conservation of 
biodiversity. Examples of natural properties 
include Sagarmatha National Park in the 
Himalayas of Nepal and Iguacu National Park, a 
cross border listing shared by Brazil and 
Argentina. 

Cultural heritage sites are equally priceless. 
Certain archaeological sites, groups of historic 
buildings, ancient towns, monumental sculptures 
and paintings whose significance transcends 
political or geographical boundaries constitute part 
of this irreplaceable heritage. Some of these 
treasures represent unique artistic achievements 
and masterpieces of the creative genius; others 



have exerted great influence over a long period of 
time or within a cultural area of the world. Some 
may be the unique witness to a civilisation which 
has disappeared or to a type of settlement that has 
become vulnerable under the impact of change. 
Others may be associated with ideas or beliefs 
that have left an indelible mark on the history of 
humanity. Cultural heritage covers a wide range 
of properties, including the Tassili n'Ajjer in 
Algeria and the Vezelay Abbey in France. 

A World Heritage Property can be listed for 
both natural and cultural values. Even so, the 
division between natural and cultural properties is 
somewhat artificial, and in the last year has been 
addressed to some extent with the addition of 
cultural landscapes as a group over the middle 
ground. 

An essential characteristic of the World 
Heritage properties is their variety: without the 
multiplicity of animal and plant species and the 
diversity of ecosystems, without the distinctive 
contributions of every culture and every people, 
the immense tapestry would remain incomplete. 

To date, Australia has ten World Heritage 
properties, which all contribute significantly to the 
wealth of the tapestry. Australian World Heritage 
properties read like a travelogue of our most 
spectacular and unique places and include: 

• the Great Barrier Reef 

• the Willandra Lakes Region 

• the Lord Howe Island Group 

• Uluru National Park 

• the Wet Tropics of Queensland 

• East coast temperate and subtropical 
Rainforest parks 

• the Tasmanian Wilderness 

• Shark Bay 

• Kakadu National Park 

• Fraser Island. 

All of Australia's properties have been listed 
for their natural values and are generally very 
large. We have the honour of having four 
properties listed on all four natural criteria, and 
three listed for both natural and cultural criteria. 
This puts our properties into a very elite group. 

Australia has been very active in the World 
Heritage field, not only with the listing of our 
properties, but also with their protection. In 1983, 
the Commonwealth Government enacted the 
World Heritage Properties Conservation Act, 
which provides for the protection and conservation 



214 



M.JINMAN 



of World Heritage Properties in Australia and its 
external territories. 

There are now 380 sites on the World Heritage 
List, which includes several fossil sites: 

• the Burgess Shale site in Canada, 

• the Dinosaur Provincial Park in Canada, 

• and Olduvai Gorge in Tanzania. 
Australia has a fossil record of great antiquity, 

extending from 3.5 billion years ago to the present. 

On 21 December 1992, the Prime Minister, Mr 
Keating announced in his Environment Statement 
that the fossil sites at Riversleigh and Naracoorte 
would be nominated to the World Heritage List in 
1993. The Murgon Fossil site will also be 
included in the nomination. Agreement has been 
reached with the South Australian and 
Queensland Governments to proceed with the 
nomination. 

The three sites represent three key stages in the 
evolution of our unique Australian fauna over the 
last 55 million years. 

The marsupial fossils from Murgon include a 
diverse suite of primitive forms, some of which 
appear to more closely resemble extinct South 
American marsupials than any previously known 
from Australia. Besides marsupials, many other 
discoveries have been made, including the tiny 
tooth of a very small placental mammal, which is 
challenging understanding about the evolutionary 
and biogeographic history of this region of the 
world. The site is about 55 million years old, and 
is the only link between the opalised monotreme 
jaw from Lightning Ridge in NSW and younger 
mammal deposits. 

The Riversleigh deposits date from 25 million 
years old to the present, and occur in unique 
freshwater limestone. With more than 20 000 
specimens representing 150 faunal assemblages, 
Riversleigh has lead to an understanding of how 
the environment and the animals that lived in it 
have changed over time from a rich rainforest 
community to a semi-arid grassland. 

The fossil bed and Ossuaries of Victoria Fossil 
Cave at Naracoorte contain the remains of at least 
93 vertebrate species, ranging from tiny frogs to 
buffalo sized marsupials. This makes it one of the 



richest Pleistocene marsupial fossil deposits in the 
world. The sediments accumulated between 170 
000 and 1 8 000 years ago and therefore over two 
of the Pleistocene glacial periods. 

As the sites occur in two different States, the 
Commonwealth Government is responsible for 
preparing the nomination documents in close 
consultation with the States involved. 
Nominations must be submitted to the UNESCO 
secretariat in Paris - by 1 October each year. They 
are then referred to the World Heritage Bureau 
(the executive of the Committee) and thoroughly 
assessed. The Bureau is assisted in this task by 
IUCN - the World Conservation Union which 
advises on natural sites and ICOMOS which 
advises on cultural sites. In addition, these 
organisations consult with relevant experts around 
the world. 

The evaluations are considered by the Bureau at 
their meeting in the year following the submission 
of the nomination, and a recommendation on the 
listing of the property is made to the World 
Heritage Committee. The Committee consists of 
21 nations and represents the different regions and 
cultures of the world. It considers the 
recommendations from the Bureau and the 
evaluation from IUCN or ICOMOS and decides 
whether the property will be inscribed. 

For the Australian Fossil Sites nomination 
being prepared this year, this timetable will mean 
the nomination is submitted by October 1993, 
assessed in early 1994, considered by the Bureau 
in June 1994 and a decision made by the World 
Heritage Committee in December 1994. As can be 
seen, World Heritage Listing is not an instant 
process and usually takes about two years to 
complete. 

This fossil sites nomination seeks to tell a story 
of change over time, rather than just providing a 
snapshot of the sites. There is potential if this 
nomination succeeds, to nominate further sites that 
fill out the story of mammal evolution in 
Australia, or tell other unique stories of universal 
value. World Heritage attracts a lot of attention, 
and the 1993 nomination of Australian Fossil 
Sites will increase worldwide awareness of our 
unique Australian history. 



Mary JINMAN, World Heritage Unit, Department of Environment, Sport and Territories G.P.O. Box 
787, Canberra, Australian Capital Territory 2601. Rec. S. Aitst. Mils. 27(2): 213-214, 1994. 



PROTECTION OF MOVABLE CULTURAL HERITAGE 

Phil Creaser 

Summary 

The Commonwealth Protection of Movable Cultural Heritage Act 1986 was enacted in 1986 and 
brought into operation in 1988. The principal goal of the program is, as stated in November 1985 in 
the Federal Parliament by the then Minister for the Arts, Heritage and the Environment, the Hon. 
Barry Cohen MP, 'to protect Australia's heritage of cultural objects and to extend certain forms of 
protection to the cultural heritage of other nations' through controls on the export and import of 
significant movable cultural heritage objects. A closely related goal was to enable Australia to 
accede to the 1970 UNESCO Convention on the Means of Prohibiting and Preventing the Illicit 
Import, Export and Transfer of Ownership of Cultural Property. 



PROTECTION OF MOVABLE CULTURAL HERITAGE 



The Commonwealth Protection of Movable 
Cultural Heritage Act 1986 was enacted in 1986 
and brought into operation in 1988. The principal 
goal of the program is, as stated in November 
1985 in the Federal Parliament by the then 
Minister for the Arts, Heritage and the 
Environment, the Hon. Barry Cohen MP, 'to 
protect Australia's heritage of cultural objects and 
to extend certain forms of protection to the cultural 
heritage of other nations' through controls on the 
export and import of significant movable cultural 
heritage objects. A closely related goal was to 
enable Australia to accede to the 1970 UNESCO 
Convention on the Means of Prohibiting and 
Preventing the Illict Import, Export and Transfer 
of Ownership of Cultural Property. 

Probably the most important aspect of the Act is 
the National Heritage Control List, which defines 
the categories of movable objects that are classed 
as 'Australian protected objects'. The Control List 
includes Class A objects which cannot be granted 
a permit for export and Class B objects that may 
be granted a permit for export. 

Class A objects include some of the most 
significant items of Aboriginal heritage: 

• bark and log coffins; 

• human remains; 

• rock art; and 

• dendroglyphs (carved burial and initiation 
trees). 

Class B objects include: 

• archaeological objects; 

• other objects of Aboriginal heritage; 

• archaeological and ethnographic objects of 
non-Australian origin; 

• natural science objects of Australian origin; 

• objects of applied science or technology; 

• military objects; 

• objects of decorative art; 

• objects of fine art; 

• books, records, documents, graphic material 
and recordings; 

• numismatic objects; 

• philatelic objects; and 

• objects of social history. 

The Control List also sets out the particular 
criteria defining 'Australian protected objects' in 
each category controlled under the Act. Generally 
the criteria include historical association, cultural 
significance to Australia, representation in an 
Australian public collection, age and current 
Australian market value. 



The Act also provides for the establishment of 
the National Cultural Heritage Committee whose 
main function is to advise the Minister on the 
operation of the Act. The Committee consists of 
ten people of whom four represent collecting 
institutions, four have experience relevant to the 
cultural heritage of Australia, one is a nominee of 
the Minister for Aboriginal Affairs and one is a 
member of the Australian Vice Chancellors 
Committee. 

The day-to-day administration of the Act is 
carried out by the Cultural Heritage Branch of the 
Department of the Arts and Administrative 
Services. The most common aspect of the 
administration of the Act is the process for the 
issuing of export permits. 

It is important to stress that within each 
category there is a definition of whether or not the 
item comes under the Act. For example, a 
palaeontological object must have an Australian 
market value of more than $1 000 before it comes 
under the Act. Similarly, a mineral must have a 
value of $10 000. However all meteorites and 
australites are covered under the Act. In many 
cases, the item does not come under the Act. If, 
however, the item is a 'protected object' it will be 
necessary to obtain the relevant form from the 
Branch for the export of the item. 

When the completed form is returned to the 
Branch, it is then sent to Expert Examiners who 
assess the application. These examiners are 
usually from collecting institutions who have a 
good knowledge of the item in question and 
similar items elsewhere in the country. The reports 
from the Expert Examiners are then forwarded to 
the Committee for consideration. The Committee, 
in turn, makes its recommendation to the 
Commonwealth Minister-at present, Senator the 
Hon. Bob McMullan-who duly decides on whether 
a permit should be granted. One important point 
to note is that the Committee does not have to 
abide by the reports from the examiners and the 
Minister does not have to accept the Committee's 
recommendation, i.e. the Minister makes the 
decision. The time taken for this process can range 
from a few days to few months, particularly if the 
item is of major significance. 

To date, over 160 applications for an export 
permit have been received by the Department and 
only one item, a painting, 'The Bath of Diana', 
has been refused an export permit. 

One other important aspect of the Act of interest 
to palaeontologists is the topic of illegal exports. 



216 



P. CREASER 



The export of fossils with a value of $1 000 5 or 
more from Australia is prohibited under the Act if 
a permit has not been obtained. The maximum 
penalty for this offence is a $100 000 fine or 
imprisonment for five years, or both. 

Following a request from this Department in 
mid-1991, the Australian Federal Police began 
inquiries into the alleged theft of Ediacaran fossils 
from South Australia. These inquiries resulted in 
the identification of a number of suspects. The 
investigation also identified further sites in South 
Australia and Western Australia where fossils had 
been removed and in late October 1991, the 
Australian Customs Service in Perth intercepted a 
suspect allegedly attempting to smuggle fossils 
out of the country. A number of fossils were seized 
which police believe were intended for sale on 
overseas markets. The police are continuing their 
inquiries and some fossils which have been 
illegally exported have been returned to Australia. 
It is not possible to provide all the details of this 
case as the investigation is still to be finalised. 8 

A review of the Act was conducted recently to 
evaluate the efficiency and effectiveness of the 
cultural heritage export and import control 
program, established by the Act and Regulations. 
The review report was released as a discussion 
paper. Government departments and authorities, 
interested organisations and the general public 
were invited to comment on the report and its 
recommendations and ninety-two submissions 
were received. The major issues addressed in the 
report included: 

• publicity for the scheme; 



• devolution of the centralised system of 
administration of the scheme to the collecting 
institutions in the States and Territories; 

• amendments to the Control List including the 
creation of a 'National Register'; 

• overcoming the lack of money in the 
National Cultural Heritage Fund. 

A total of 60 recommendations were made in 
the report, some of which require changes to the 
Act. Of particular interest to palaeontologists is 
the recommendation to remove the $1 000 
minimum market value on palaeontological 
objects. If accepted, this will mean that all fossils 
will come under the ambit of the Act. A report 
will be presented very shortly to the Minister 
which addresses these recommendations in terms 
of the public response, the views of the Committee 
and the department perspective. Depending on 
ministerial decisions, there may be some 
amendments to the Act and some changes to the 
administration of the scheme. 

Movable cultural heritage is a complex topic. 
There is a need for the Act to be administered in a 
very understanding way and for policies to be 
developed which reflect current trends and 
financial positions. However, it could be 
considered that awareness of the scheme by 
individuals as well as collecting institutions is the 
most important issue that needs to be addressed. 
In this regard, a mailing list is being compiled of 
all those with an interest in this topic. 

With an increasing awareness of the scheme, 
we are confident that there will be an increasing 
awareness of the value of our cultural heritage, 
including our palaeontological heritage. 



Phil CREASER, Cultural Heritage Branch, Department of the Arts and Administrative Services, G.P.O. 
Box 1920, Canberra, Australian Capital Territory 2601. Rec. S. Aust. Mus. 27(2); 215-216. 1994. 



§ On 3 August 1993, the Regulations under the Act were amended and the $1 000 minumum value for 
palaeontological objects was removed. This means that a permit is required before any fossil secimen is exported. 
Following a Cabinet reshuffle on 28 January 1994, the Hon. Michael Lee, Minister for Communications and the Arts, 
assumed responsibility for the Act and administration of the scheme. At that time, the number of applications for 
permits had increased to more than 220 with the majority of recent applications being for fossils. In addition, one 
further item, an historic steam engine, has been refused an export permit. 



ABSTRACTS OF THE FOURTH CONFERENCE ON AUSTRALIAN 
VERTEBRATE EVOLUTION, PALAEONTOLOGY AND SYSTEMATICA 

ADELAIDE, 19-21 APRIL, 1993 



ARCHER, M., GODTHELP, H., MUIRHEAD, J., NOCK, C, & AUGEE, M. 1994 
School of Biological Sciences, University of New South Wales, Kensington, 

New South Wales, 2033. 



Summary 



Discovery in late 1990 of a single upper molar of an early Paleocene monotreme by 
palaeontologists from the University of La Plata (including Rosendo Pascual and Edgardo Ortiz 
Jaureguizar) led to the description of Monotrematum sudamericanum (Pascual et al. 1992). An 
invitation from the Argentinians to mount a joint expedition from the University of New South 
Wales was taken up with the aid of a substantial financial grant from the Australian Geographic 
Society and equipment donated by Paddy Pallins of Sydney. In November and December 1992, we 
combined efforts to investigate early Paleocene Banco Negro Inferior exposures at Punto Pellegro 
north of Comodoro Rivadavia, Argentina. The results were highly successful. In addition to more 
ornithorhynchid material, marsupials, condylarths, engimatic mammals and colossal leptodactylid 
frogs were found on eroded surfaces and in quarries. Exploration of late Mesozoic to Oligocene 
sediments in the Grand Baranca, near Sarmiento (west of Rivadavia Comodoro) was equally 
productive and provides, with the Punto Pellegro deposits, a significant opportunity to expand 
understanding about the earliest Cainozoic mammal record of South America. 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 217 

ABSTRACTS OF THE FOURTH CONFERENCE ON AUSTRALASIAN VERTEBRATE 

EVOLUTION, PALAEONTOLOGY AND SYSTEM A TICS, 

ADELAIDE, 19-21 APRIL 1993 



In pursuit of the peregrinating Patagonian platypus 

ARCHER, M, GODTHELP, H., MUIRHEAD, J., NOCK, C. & AUGEE, M. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

Discovery in late 1990 of a single upper molar of an early Paleocene monotreme by palaeontologists 
from the University of La Plata (including Rosendo Pascual and Edgardo Ortiz Jaureguizar) led to the 
description of Monotrematum sudamericanum (Pascual et al. 1992). An invitation from the 
Argentinians to mount a joint expedition from the University of New South Wales was taken up with 
the aid of a substantial financial grant from the Australian Geographic Society and equipment donated 
by Paddy Pallins of Sydney. In November and December 1992, we combined efforts to investigate early 
Paleocene Banco Negro Inferior exposures at Punto Pellegro north of Comodoro Rivadavia, Argentina. 
The results were highly successful. In addition to more ornithorhynchid material, marsupials, 
condylarths, engimatic mammals and colossal leptodactylid frogs were found on eroded surfaces and in 
quarries. Exploration of late Mesozoic to Oligocene sediments in the Grand Baranca, near Sarmiento 
(west of Rivadavia Comodoro) was equally productive and provides, with the Punto Pellegro deposits, a 
significant opportunity to expand understanding about the earliest Cainozoic mammal record of South 
America. 



Australian VP bibliography computer data base 

BAYNES, A. 1994 

Department of Earth and Planetary Sciences, Western Australian Museum, Perth, Western Australia 

6000. 

The first version of the VP bibliography computer data base was created by combining the Australian 
mammal chapter bibliography and appendix (in which mammal genera are indexed to the papers) from 
the then-unpublished Vertebrate Palaeontology of Australasia. It is available in Macintosh and PC 
versions, and its purpose is to answer the question 'What is the literature on genus xT It was offered 
free on a flyer in VP of Australasia. The response was, frankly, lousy. Undismayed, I am expanding it 
into the MARB (Mammals Amphibia Reptiles & Birds) Australian VP bibliographic data base. In 
MARB both the process of extracting references to a mammal genus and the indexing of mammal 
genera mentioned in a paper are simpler. Such a data base can potentially be perfected for the older 
references with complete generic indexing, and the references can always be cited without error. It can 
also be kept up-to-date much more easily than a reference book. By CAVEPS, MARB will include the 
papers but not the indexing of the ARB genera. For these expert help is needed. Graduate students 
probably represent the best source of manpower for completing the indexing and adding new references, 
as they will be making the greatest use of the data base. I am hoping someone else will do one on fossil 
fish, which I know nothing about. 

The MARB data base was demonstrated at CAVEPS 93. Copies are available: free to those who 
contribute to it; $10.00 (or $1 1 .00 including the disk) for others. 



218 ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 

Preliminary anaylsis of mammals from Aliens Cave, southern Nullarbor 

BAYNES', A. & WALSHE 2 , K. 1994 

1. Department of Earth and Planetary Sciences, Western Australian Museum, Perth, Western Australia 
6000. 

2. Department of Archaeology and Anthropology, The Faculties, Australian National University, 
Canberra, Australian Capital Territory 2601 . 

Archaeological excavations made in Aliens Cave in 1989 yielded substantial quantities of bones. 
Those from pit E4 (1 m 2 ) have been analysed. The stratigraphy consists of a basal orange unit 1.9 m 
deep terminating in a gravel lag deposit that marks a disconformity; above this is a 1 .4 m dark grey unit. 
Two radiocarbon dates have been obtained: 3 720 ± 150 yrs BP at about 1.1m depth and 5 860 ± 430 
yrs BP at the base of the dark grey unit. The orange unit is currently undated, but probably late 
Pleistocene in age. Remains of large mammals are generally highly fragmented; those of the smaller 
mammals are dissociated but more complete. Burnt bone is present at all levels. These observations 
suggest that the principle agents of accumulation were owls (Tyto spp.), devils and humans. In the 
orange unit the contribution from devils and humans predominated; in the dark-grey unit there is more 
substantial owl prey component. 

The mammal fauna of the orange unit is restricted to species whose original distributions include the 
most arid parts of modern Australia. Above the disconformity species richness increases with the 
addition of some arboreal species and a substantial south-western element, but few losses. These faunas 
suggest very arid conditions at the time of deposition of the orange unit and higher rainfall during 
deposition of the dark grey unit. This is consistent with vegetation changes inferred from palynological 
investigations of this and other southern Nullarbor deposits. 



Responses of mammalian communities to late Quaternary climatic changes 

BAYNES 1 , A. & WELLS 2 , R. T. 1994 

1. Department of Earth and Planetary Sciences, Western Australian Museum, Perth, Western Australia 
6000. 

2. School of Biological Sciences, Flinders University of South Australia, Bedford Park, South Australia 
5042. 

Compared to North America, Australia still has very few windows opened on what was happening to 
mammalian communities in the geologically recent past. The principal southern and western sites are 
Devils Lair and Skull, Hastings, Aliens, and Victoria Caves. All analysed large stratigraphically discrete 
samples of mammals show log-normally distributed rank-abundance distributions, indicating that a 
number of communities has been sampled by agents accumulating remains in the caves, as would be 
expected. In only one or two cases has an attempt been made to resolve the mammals from a deposit 
into their contemporary communities; the other sites merely provide assemblage data. The small to 
medium-sized mammal assemblages from the sites reflect different species equilibria in local mammal 
communities at different times in the glacial-interglacial cycle. The sites in temperate and semi-arid 
areas show greater species richness in glacial age deposits than in later Holocene deposit the so-called 
disharmonious assemblages effect. In marked contrast, in Aliens Cave, located on the edge of the 
modern arid zone but close to the Southern Ocean coast, the higher species richness is in the Holocene 
levels. This pattern is consistent with the idea that species richness is higher at times of more equable 
climates. As climates become more equable increasing numbers of species appear to have been 
integrated into more complex communities. As climates changed in the opposite direction faunal 
segregation has resulted in simpler communities. The general pattern is also consistent with a centrifugal 
movement of faunas in response to extremes of continental glacial aridity. 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 219 

New species of Nambaroo (Flannery and Rich) from Riversleigh, northwestern Queensland. 

COOKE, B.N. 1994 

School of Life Science, Queensland University of Technology, Brisbane, Queensland, 4001 . 

New species of Nambaroo (Flannery and Rich, 1986) are described from a variety of sites of 
estimated early to mid-Miocene age from Riversleigh deposits of northwestern Queensland. The new 
species are phenetically similar to those described by Flannery and Rich (1986) from the mid-Miocene 
Namba Formation of South Australia. Their lower molar morphology supports the views of those 
authors regarding the evolution of the hypolophid and posterior cingulum among macropodids. 



Pliocene whales and dolphins (Cetacea) from the Vestfold Hills, Antarctica 

FORDYCE 1 , R. E. & QUILTY 2 , P. G. 1994 

1 . University of Otago, Dunedin, New Zealand. 

2. Australian Antarctic Division, Kingston, Tasmania, 7050. 

Early Pliocene cetaceans from Marine Plain, Vestfold Hills (68°35'S, 78°00'E), are the only fossil 
higher vertebrates known so far from the Antarctic Oligocene-Pleistocene. Of these, one (possibly two) 
undescribed extinct species of dolphin (Odontoceti: Delphinidae: new genus) is known from skulls, 
mandibles, ear bones, and post-cranial elements from three individuals. Skull bones have contact 
relationships similar to those in other delphinids, but skull topography is remarkably convergent with 
some extant toothless beaked whales in the genus Mesoplodon (Odontoceti: Ziphiidae): the narrow 
rostrum is toothless, there is a prenarial basin, and the premaxillae carry flat vertical flanges beside the 
nares. The dolphins seems too derived to be ancestral to any extant species; otherwise, relationships to 
extant delphinids are uncertain. The shallow marine depositional setting at Marine Plain cautions 
against but does not preclude ziphiid-like habits (pelagic, deep-diving, semi-solitary, squid-eating) for 
the dolphin. 

Of other specimens, an articulated series of large vertebrae is probably from a baleen whale 
(Mysticeti). A small right whale (Mysticeti: Balaenidae: genus and species uncertain) has a narrow and 
ventrally curved but otherwise poorly preserved rostrum and a fragmentary braincase. Ear bones from 
this specimen, when fully prepared, should better reveal relationships. The fossils are unexpected in that 
small dolphins (or small ziphiids) and small right whales appear ecologically insignificant in near-shore 
modern Antarctic waters. Indeed, late Neogene Cetacea elsewhere include bizarre forms and/or unusual 
distributions, as well as species and distributions of modern aspect. These unusual taxonomic and 
ecological patterns apparently did not persist into the Pleistocene, which suggests a later Pliocene shift 
in global cetacean ecology. 



Cuddie Springs: new light on Pleistocene megafauna 

FURBY 1 , J. & JONES 2 , R. 1994 

1. School of Geography, University of New South Wales, Kensington, New South Wales, 2033. 

2. Department of Palaeontology, Australian Museum, Sydney, New South Wales, 2000. 

The faunal list for the Cuddie Springs site (compiled after excavations in 1993) has been expanded to 
include an additional 11 taxa following excavations during 1991. Pollen and sediment analysis has 
provided a palaeoenvironmental record spanning the late Pleistocene, including the period when the 
megafauna disappeared. The vegetation history from Cuddie Springs has a time depth not previously 
recorded from arid-semi-arid contexts. Archaeological material at depth was first documented in the 
recent excavations with aboriginal stone artefacts occurring in direct association with bones of 
Genyornis, Diprotodon and Sthenurus, dating to about 30 000 years B.P. The faunal record is 
continuous to at least five metres depth and one of the aims of ongoing research is to document the 
changes in the faunal assemblage through time in an environmental context. 



220 ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 

Some rodents from the Pleistocene fluviatile deposits of the eastern Darling Downs, Queensland 

GODTHELP,H. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

Sediments collected from several sites on the Eastern Darling Downs were washed and sorted. The 
fossilised remains of many vertebrates were recovered. The assemblages were dominated by the remains 
of murid rodents; 8 taxa are recognised. One taxon is as yet unidentified and may represent a new 
species of Pseudomys, the remaining are all extant species. Mastacomys fuscus and Pseudomys higginsi 
are recorded from Queensland for the first time. 



A new species of the murid Zyzomys from the Pliocene Rackhams Roost Deposit, northwest 
Queensland 

GODTHELP,H. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

A new species of the murid Zyzomys is described on the basis of fossil material from Rackhams 
Roost, a Pliocene Macroderma roost site on Riversleigh Station. This species is dentally the most 
primitive of all known species of rockrats. It is the most abundant mammal species in the deposit. There 
is a similarly high abundance of Zyzomys species in more recent Macroderma deposits. 



Temporal fenestration in a procolophonid reptile 

HAMLEY, T. & THULBORN, T. 1994 

Department of Zoology, University of Queensland, Queensland, 4072 

The Late Permian and Triassic procolophonids were late survivors from the initial radiation of 
amniotes, and their skulls are usually thought to have retained the primitive anapsid condition, without 
temporal openings. However, two skulls attributed to Procolophon trigoniceps, from the Lower Triassic 
of South Africa, have been found to possess a lower temporal opening resembling that of the synapsid 
reptiles. The temporal opening was first discovered by H. G. Seeley 120 years ago, but its existence has 
been overlooked or denied in palaeontological literature for the past 90 years. Rediscovery of the 
temporal opening prompts us to examine the possible affinities of procolophonids and other 
'parareptiles'. In doing so, we find no substantial support for the recent suggestion that procolophonids 
are the sister group of turtles. 



Morphological changes in the Australian Macroderma lineage (Microchiroptera: 
Megadermatidae) from the Oligo-Miocene to the present 

HAND, S.J. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

From the Riversleigh Tertiary deposits, eight megadermatids belonging to two lineages are now 
known. Morphological changes in the Australian Macroderma lineage are traced from the Oligo- 
Miocene to the present and comments are made on broader aspects of megadermatid evolution. In some 
Riversleigh deposits, there is evidence for sympatry of very differently-sized megadermatids. The 
Riversleigh megadermatids have provided an opportunity to trace an apparent trend to shorten the face 
in the Macroderma lineage from the Oligo-Miocene to the present and to examine a tendency to 
gigantism in independent megadermatid lineages. 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 221 

A new and distinctive Oligo-Miocene hipposiderid (Microchiroptera: Hipposideridae) from 
Riversleigh Station, Queensland 

HAND, S.J. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

A large, new and very distinctive hipposiderid is described from complete skull material recovered 
from the Oligo-Miocene Bitesantennary cave deposit located on the south-eastern margin of the D-Site 
Plateau, Riversleigh Station, north-western Queensland. The species is compared with Tertiary French 
and Australian species of Brachipposideros as well as Quaternary Australian hipposiderids (genera 
Rhinonicteris and Hipposideros). The phylogenetic relationships of the new bat and apparent problems 
in current hipposiderid taxonomy are discussed. 

Scaly-foots and pointy teeth: pygopod mandibular variation and first fossil record from the 
Miocene of Queensland 

HUTCHINSON, M. N. 1994 

South Australian Museum, Adelaide, South Australia, 5000. 

The snake-like pygopod lizards are a group of 35 species confined to Australia and New Guinea. 
They are gekkonoid, but their phylogenetic relationship to typical geckoes is not yet well established. 
Recent morphological and emerging biochemical data point to a sister group relationship with, or 
possibly within, the Australian diplodactyline geckoes. Pygopods show considerable variation in 
mandibular and dental structure, correlated with dietary specialisation in several genera. This variation 
is summarised and employed in identifying the first fossil member of the family to be reported, a partial 
dentary, from the early Miocene of the Riversleigh area, northwestern Queensland, Australia. The fossil 
is most similar to the living genus Pygopus but differs in its more evenly-sized teeth. 

The Neogene radiation of Australian sharks: real or apparent ? 

KEMP,N.R. 1994 

Tasmanian Museum, Hobart, Tasmania, 7000. 

By the Mid-Tertiary all eight orders of Neoselachians - modern sharks - were represented in 
Australia. Numerically, the Lamniformes (mackerel sharks) dominate the Tertiary record. The 
Palaeogene is well represented by the Odontaspididae and the Mitsukurinidae, by Carcharias (grey 
nurse sharks) and Scapanorhynchus (goblin sharks), respectively. The Odontaspididae continue through 
to the Recent with large numbers of teeth being preserved, especially in the Neogene. However, the 
Lamnidae dominate the Neogene in the number of species, with up to a dozen taxa representing at least 
four genera: Carcharodon (white pointers), Carcharoides (serrated porbeagles), hums (makos) and 
Lamna (porbeagle). The presence of at least seven species of hums in Australia in the Neogene may be 
indicative of a southern centre of radiation for this genus; nowhere else do all these species occur 
together. The Carcharhiniformes (ground sharks) which today contain, world-wide, more than half of all 
shark species are relatively poorly represented in Australian Neogene deposits, and are conspicuous by 
their virtual absence from the Palaeogene. Of this order, Carcharhinus (whalers) is the most common 
taxon, followed by Galeocerdo (tiger shark), Galeorhinus (school shark), Hemipristis (snaggletooth 
shark), and Sphyrna (hammerhead sharks) which is known from only a few teeth. The Hexanchiformes 
(six- and sevengill sharks), Pristiophoriformes (saw sharks), Heterodonti formes (bull sharks) and 
Orectolobiformes (wobbegongs) are known variously throughout the Tertiary, but are certainly not 
common. The Squaliformes (dogfish sharks) have yet to be found. The influence of the warmer Miocene 
temperatures - which would be expected to result in an increase in the number and variety of sharks in 
southern Australian seas is not reflected in the local fossil record. The presence in the Australian 
Neogene of many species of hums is real, the numerical dominance of /. hastalis is due to a collecting 
bias, Carcharias is definitely a common taxon. The relative paucity of most other shark taxa may be due 
to depositional features such as fossiliferous winnowed deposits, and lack of preservation of certain 
environments of deposition. 



222 ABSTRACTS; CAVEPS 1993. REC. S. AUST. MUS. 27(2) 

Studies of the Late Cainozoic diprotodontid marsupials of Australia 1. Revision of the genus 

Euowenia 

MACKNESS,B. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

Species of the genus Euowenia are rare taxa in Australian faunal assemblages. Two species have 
been described: Euowenia grata (De Vis 1887) on the basis of a partial cranium and dentary from 
Chinchilla, Queensland and E. robusta (De Vis 1891) from dentaries found at Freestone Creek, near 
Warwick, Queensland. Woods (1968) suggested that E. robusta was a junior synonym of Nototherium 
inerme. Mackness (1988) identified E. robusta as a zygomaturine and this taxonomic assessment is 
confirmed here. A number of additional specimens of Euowenia have been recovered from northern 
Australia including a new species, described here on the basis of a partial cranium and lower dentary 
featuring relatively unworn teeth. This additional material has highlighted a number of useful 
synapomorphies for Euowenia and a new generic diagnosis is provided. Additional published records of 
Euowenia are examined in the light of this new diagnosis. A significant extension in range and 
chronology is reported. A guide to identifying isolated Euowenia teeth from those of other northern 
Australian diprotodontids is also provided. The late Miocene Meniscolophus mawsoni is shown to 
share many synapomorphies with the genus Euowenia and a new taxonomic position is proposed for 
this taxon. 



Studies of the Late Cainozoic diprotodontid marsupials of Australia 2. The identity of 
Zygomaturus macleayi Krefft and Z. creedii Krefft 

MACKNESS, B. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

A partially reconstructed palate with left and right PMVF was described from the Condamine River, 
Queensland by Krefft in Longman (1921) as Zygomaturus macleayi. Longman recognised that the P 3 of 
Z. macleayi was not the same as Z trilobus but could not make any definitive taxonomic assignment. It 
is confirmed here that the holotype does not represent a zygomaturine based on its P 3 morphology. 
Another right P 3 -M 5 , figured by Owen (1872) as Nototherium inerme and a cast of a left P 3 -M 5 
(BMNH M5002 labelled Nototherium mitchelli although presumably from the same animal) represent 
additional specimens referable to Zygomaturus macleayi. Woods (1968) suggested that the 
identification of the Owen illustration as Nototherium inerme was correct. This cannot be the case, 
however, as the molar row gradient of Zygomaturus macleayi is substantially different to all 
Nototherium holotypes and lectotypes, even allowing for allometric distortion. Zygomaturus macleayi 
cannot be placed in any existing diprotodontid genus although it does share many features in common 
with the Pliocene Euryzygoma. It is therefore placed in a new monotypic genus. 

Zygomaturus creedii was described by Krefft (1873) on the basis of a fractured right premaxilla with 
I 1 and other incisor fragments. Only the incisor remains in the Australian Museum. The holotype is 
indistinguishable from other Fs from Zygomaturus trilobus and therefore Z creedii is regarded as its 
junior synonym. 



Studies of the Late Cainozoic diprotodontid marsupials of Australia 3. Key to the identification of 
lower molars 

MACKNESS, B. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

The fossil remains of diprotodontids are commonly found as isolated molars or as dentary and/or 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 223 

maxillary fragments. A lack of recognisable genera-specific features has resulted in confusing taxonomic 
assignments. Examination of an extensive collection of late Cainozoic dentary fragments from Australia 
has revealed a number of features useful for distinguishing genera. The poster presents a key to the 
identification of the lower molars of Diprotodon, Euowenia, Euryzygoma, Nototherium and 
Zygomaturus. This, along with a guide to the lower premolars presently in preparation by Mackness 
should provide the basis for more consistent determination of diprotodontid species diversity and 
distribution in the late Cainozoic of Australia. 



An enigmatic family of marsupials from the Early Pliocene Bluff Downs Local Fauna of 
northeastern Queensland 

MACKNESS, B., ARCHER, M. & MUIRHEAD, J. 1994 

School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 2033. 

Discovery of a right mandible from the early Pliocene Bluff Downs Local Fauna of the Allingham 
Formation, northeastern Queensland, provides evidence for a highly specialised group of marsupials 
that has no previously known representatives. Key features are tribosphenid-like molars with elaborate 
buccal and anterior cingula, unique buccal cusps, complex cusp and crest relationships in a gradient 
that increases from M to M 2 , complex P morphology including an elaborate postenior cusp and a 
prominently bicuspid P 2 , The M and M 4 have some features reminiscent of peramelemorphians but 
none of these are undoubted synapomorphies. The M 2 and premolars exhibit other features that are 
diprotodontian-like although these may be convergent. Some aspects of lower morphology most closely 
resemble those seen in late Cretaceous glasbiids from North America and early Tertiary didelphimorphs 
(possibly glasbiids) from South America. These resemblances too are most likely to be convergent. 
Precise occlusal and possibly thegotic wear on all blades indicates that this extraordinary morphology is 
matched by precise occlusal counterparts in the as yet unknown upper dentition rather than abnormal 
and that this individual lived for a long time. Because there are no undoubted synapomorphies shared 
with any previously known family, we propose a new monotypic family for this taxon. Its highly 
distinctive morphology suggests antiquity that well predates the early Pliocene and demonstrates that 
not all contemporary lineages of middle Tertiary marsupials were sampled by the otherwise diverse 
Oligo-Miocene communities of South Australia and Queensland. 



The Spring Park Local Fauna, a new Late Tertiary fossil assemblage from northern Australia 

MACKNESS 1 , B., MCNAMARA 2 , G., MICHNA 3 , P., COLEMAN 3 , S. & GODTHELP 1 , H. 1994 

1. School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 
2033. 

2. Department of Geology, James Cook University, Townsville, Queensland, 481 1 . 

3. Townsville Grammar School, Townsville, Queensland, 4811 . 

Preliminary investigation of a series of complex freshwater, fluviatile quartz sand and clay deposits 
from Blaggard Creek, northwest of Charters Towers, northeastern Queensland has revealed a rich 
collection of fossils. The fauna, here named the Spring Park Local Fauna, contains: Zygomaturus sp. cf. 
Z trilobus , a new species of Euowenia, Palorchestes parvus, another palorchestid, a phascolarctid, 
several macropodids, Pallimnarchus sp. cf. P. pollens, a new crocodilian and a python as well as 
remains of turtles and fish. The deposit is subjacent to Pliocene basalts. Sediment grain size and 
sedimentary structures mitigate against the idea that the deposit may have resulted from accumulation at 
the edge of a basalt bluff and suggests a stream too large to have been in this position after basalt 
deposition. Therefore, the deposit is nominally assessed as being of Pliocene age based on its 
geomorphology and biocorrelation with Chinchilla and Bluff Downs faunas. 



224 ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 

Three-dimensional analysis of variation in marsupial teeth using computed tomographic scans 

MACKNESS 1 , B. & SELDON 2 , L. 1994 

1. School of Biological Sciences, University of New South Wales, Kensington, New South Wales, 
2033. 

2. Department of Otolaryngology, University of Melbourne, Parkville, Victoria, 3052. 

The analysis of variation in the diagnostic P 3 of Zygomaturus trilobus has been undertaken by means 
of computed tomography. The study was carried out in conjunction with the Cochlear Implant Program 
of the Department of Otolaryngology, University of Melbourne, which uses computed tomographic 
scans for reconstructing intricate anatomical features such as the temporal bone to assist surgeons. The 
input data was obtained via a series of CT scans taken at 1.00 mm intervals, from the alveolus to the 
tooth crown, on a suite of isolated P 3 teeth suspended in a polystyrene foam collar. The scans, on film, 
were then digitised from a light box using a Panasonic WV-BL200 video camera connected to a IBM 
AT compatible personal computer and stored in a CompuServe GIF graphics interchange format. A 
Data Translation DT2851 medium resolution video card with 512 x 512 pixels and 256 colours was the 
only 'special' piece of hardware used in the study. An automatic edge detection system was used to 
combine the scanned images to form a 3D computer model. Automatic analysis of a number of variables 
was then possible. This included plotting the ratio of the length and width along the entire midline of 
the tooth at 1.00 mm intervals. Individual teeth were also divided into four quadrants and analysed for 
shape using the formula (10-4071A / P 2 ) as well as for volume and surface area. The study highlights the 
effectiveness of computers in assessing previously subjective characters such as 'shape', surface area 
and quadrant volumes as well as providing a suitable technique for close examination of variation in a 
number of isolated individuals. 



The morphology and relationships of thelodonts, Siluro-Devonian agnathans 

MARSS 1 , T. & RITCHIE 2 , A. 1994 

1. Institute of Geology, Tallinn, Estonia 

2. Australian Museum, Sydney, New South Wales, 2000. 

Thelodonts are a group of poorly-known Siluro-Devonian agnathans characterised by a micromeric 
dermal skeleton consisting entirely of small scales. Complete thelodonts are very rare but articulated 
specimens are known from sites in Scotland, England, Estonia, Norway and Canada. Isolated scales, by 
contrast, are often locally abundant, widely distributed in Siluro-Devonian sediments and are being 
increasingly used in biostratigraphic studies world-wide, including Australia and Antarctica. Recent 
discoveries of articulated thelodonts provide new information on the variation in shape of thelodont 
scales in different parts of the dermal skeleton, the position of the eyes, the nature of the mouth and the 
branchial apparatus, the size and the shape of the lateral, dorsal, ventral and caudal fins. The 
relationships of thelodonts with other Palaeozoic agnathans will be discussed. 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 225 

The Late Miocene Ongeva Local Fauna from the Waite Formation of central Australia 

MEGIRIAN 1 , D., MURRAY 1 , P. F. & WELLS 2 , R. T. 1994 

1 . Northern Territory Museum, Darwin, Northern Territory, 0800. 

2. Flinders University of South Australia, Bedford Park, South Australia, 5042. 

The Ongeva Local Fauna (LF) from the Waite Formation of central Australia is biochronologically 
significant because it is in litho strati graphic superposition to, and unconformably separated from, the 
Alcoota LF. It contains a zygomaturine diprotodontid (description in press) that is structurally 
intermediate between Kolopsis torus Woodburne from the Alcoota Local Fauna, and Zygomaturus gilli 
Stirton from the Beaumaris LF of Victoria. Kolopsis torus is also present in the Ongeva LF, though the 
Ongeva LF specimens differ slightly in some measurements from the Alcoota population. A 
morphospecies distinction of the two populations is not justified on the available evidence. The 
indications are that a cladogenetic event occurred in the Zygomaturinae between Alcoota and Ongeva 
LF times, an interpretation consistent with the presence of a Zygomaturus sp. and a Kolopsis sp. in the 
younger Beaumaris LF. Dromornithidae are also common elements in the Ongeva LF, but have not yet 
been analysed in detail. The indications are that a Dromornis sp. cf. D. stirtoni and an llbandornis sp. 
are present. A crocodilian, Quinkana sp., is represented by a dentary and a relatively large number of 
ziphodont teeth. The most remarkable fossils, however, are large, loosely-deposited coprolites draped 
over the bones. Two quarries are now producing Ongeva LF material, but their yields may eventually be 
limited by the huge overburdens that accompany excavating into the sides of steep hills. 



Late Cainozoic crocodilians collected by the 1980 and 1983 FUAM Expeditions into the Lake 
Eyre Basin of South Australia 

MEGIRIAN 1 , D., WELLS 2 , R. T. & TEDFORD 3 , R. H. 1994 

1. Northern Territory Museum, Darwin, Northern Territory, 0800. 

2. Flinders University of South Australia, Bedford Park, South Australia, 5042. 

3. Department of Vertebrate Paleontology, American Museum of Natural History, New York, N. Y., 
10024, United States of America. 

The Flinders University-American Museum of Natural History (FUAM) collection of crocodilian 
material from Pliocene and Pleistocene strata of the Lake Eyre Basin is just one of several collections 
made since 1892. Virtually none of the material of this geological age and provenance has been 
described, but has been variously assigned to the extinct form Pallimnarchus pollens de Vis, or to the 
extant species Crocodylus porosus Schneider. The distinction of fragmentary remains of these two taxa 
has proved difficult in the past. Poor stratigraphic control has also hampered the interpretation of 
crocodilian succession. The stratigraphy of the Lake Eyre Basin Pliocene and Pleistocene has now been 
resolved in some detail, and the stratigraphic provenance of the FUAM material is recorded. There is 
prima facie evidence in the FUAM collection, from specimens that could not have been re-worked, for 
the presence of a Pallimnarchus-hke form in the mid Pliocene (estimated age: 3.9-3.4 Ma) Tirari 
Formation and the late Pleistocene (estimated age, 0.2 Ma) Kutjitara Formation. There is no evidence in 
the FUAM collection for the presence of a Crocodylus-Yike form in the Tirari Formation. One 
Crocodylus-\\ke specimen from the Kutjitara Formation differs in preservation from other material of 
that provenance and is possibly reworked. Only two potentially diagnostic specimens were collected 
from the latest Pleistocene (estimated age: 0.04 Ma) Katipiri Formation: a jugal that may be referable to 
Crocodylus, and a maxillary fragment most closely resembling Australosuchus clarkei Willis and 
Molnar from the mid-Tertiary Etadunna Formation which forms the basement to the Pliocene and 
Pleistocene strata. 



226 ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 

Post-cranial descriptions of Ilaria and Ngapakaldia (Vombatiformes, Marsupiala) and the 
phyiogeny of the vombatiforms based on post-cranial morphology 

MUNSON,C. 1994 

Moreno Valley, CA 92553, United States of America. 

The post-crania of the vombatiform marsupial Ilaria illumidens from medial Miocene strata of South 
Australia are described and compared to those of other vombatiforms, with the observation that Ilaria 
shares a similar morphology of the manus and pes with living wombats. While this indicates a certain 
degree of fossorial activity, the size and vertebral morphology of Ilaria argue against a burrowing 
lifestyle. Another medial Miocene vombatiform, Ngapakaldia tedfordi, is described as having a 
plesiomorphic vombatiform skeleton similar in many ways to that of the phalangeriform possums, but 
with adaptations for greater size and a plantigrade, terrestrial habitus. Besides stouter and more robust 
limbs, these adaptations are evident in the concave dorsal surface and laterally facing fibular facet of the 
astragalus that creates a less flexible upper ankle joint. 

For this study, a cladistic analysis was made using the post-crania of all the families in the 
Vombatiformes and several species representing outgroups, in order to establish synapomorphies uniting 
the group and to evaluate the position of these two genera within it. The results indicate that the ilariids 
and vombatids probably share a common ancestor, based on the similarity of the metapodials and 
phalanges, especially the uniquely identical morphology of the proximal metapodial facets. 
Ngapakaldia 's similarity in form to phalangeriform possums reflects the arboreal ancestry of the 
vombatiform clade and indicates the plesiomorphic state from which the post-crania of other, more 
specialised vombatiform families (i.e. fossorial wombats and ilariids) are derived. 



Recent Diprotodon discoveries in South Australia 

PLEDGE, N. S. 1994 

South Australian Museum, Adelaide, South Australia, 5000. 

From the early days of the colony, Diprotodon remains have been found. Prior to the Lake Callabonna 
discoveries, the best were a skull from near Gawler in 1891 (now apparently lost) and a skull from 
Baldina Creek near Burra in 1890. Since 1970, and the publicity engendered by the expedition to Lake 
Callabonna in that year, four chance discoveries have been made in the Adelaide area, and two on the 
West Coast. More material has been obtained from a Port Pirie site, and minor records made for the 
Woomera area, Morgan and Naracoorte. 

A badly tumbled and rolled molar found in beach gravels at Hallett Cove in 1970 may now be related 
to a partial pelvis and other bones and teeth found in 1992 in the banks of the Field River which flows 
into the southern end of the cove. Soil samples from this site have been taken for thermoluminescent 
dating by Prof. J. Prescott, University of Adelaide. A similar discovery was made in early 1993 in the 
Little Para River, near Salisbury. In 1980, a jaw fragment was found in a sewer trench near Darley 
Road, Paradise, on the River Torrens. Diprotodon jaws and other bones have been found in lithified 
Bridgewater Formation (coastal dune sand) at Sheringa Beach, west of Port Lincoln, and also near Head 
of the Bight, west of Ceduna. These bones are well preserved but firmly embedded in the rock. Jaws and 
a partial skeleton have been recovered from an old locality - a sand quarry near Port Pirie, but have been 
damaged by the bulldozers. A poorly preserved skeleton is known in Elizabeth Creek which flows into 
Pernatty Lagoon near Woomera. 

Mineralised tooth fragments have been collected from river alluvium excavated from a farm dam near 
Morgan, on the River Murray, and several teeth were found during excavations in Henschke Fossil 
Cave at Naracoorte. Bones embedded in reef rock at a depth of five metres off Victor Harbor have been 
reported but not yet confirmed. 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 227 

A new extinct sthenurine kangaroo (Marsupiala, Macropodidae) from southeastern Australia 

PRIDEAUX, G. J. & WELLS, R. T. 1994 

School of Biological Sciences, Flinders University of South Australia, Bedford Park, South Australia, 

5042. 

A new species of Simosthenurus is described from Late Pleistocene deposits in southeastern South 
Australia, western Victoria, northern and southeastern New South Wales, northwestern Tasmania and 
the Nullarbor Plain, Western Australia. The species is similar in size and mandibular morphology to 
Simo. brownei, but its cranium is more dolichocephalic with a less inflated nasal region. The dentition 
is distinctive and, although similar in size to Simo. occidentalism several molar characters appear closer 
to Sthenurus andersoni in morphology. Because the new species exhibits features previously only 
considered typical of either Simosthenurus or Sthenurus, a thorough reassessment of sthenurine generic 
relations is impelled. Whether or not the distribution of craniodental character states within the groups 
confirms the tentative placement of this new form within Simosthenurus \ it probably evolved soon after 
split of the Simosthenurus and Sthenurus lineages. 

An ornithomimosaur and protoceratopsian from the early Cretaceous of southeastern Australia 

RICH 1 , T. H. & VICKERS-RICH 2 , P. 1994 

1 . Department of Paleontology, Museum of Victoria, Melbourne, Victoria, 3000. 

2. Department of Earth Sciences, Monash University, Clayton, Victoria, 3168. 

Ornithomimosaurs were intermediate-sized, agile, edentulous theropod dinosaurs best known from 
the late Cretaceous of the Northern Hemisphere. Fragmentary records of them exist there in the early 
Cretaceous and possibly the late Jurassic. Elaphrosaurus bambergi from the late Jurassic Tendaguru of 
Tanzania is likely to be a primitive member of this group of dinosaurs. 

A pair of femora, one of an adult and one of a juvenile about 45% as large as the former were found 
at the late Aptian - early Albian Dinosaur Cove site, Victoria, Australia, in 1991. The relatively large 
size of the lateral distal condyle in comparison to the medial distal condyle characterises these specimens 
and the ornithomimosaurs among theropods (Barsbold & Osmolska 1990). The lesser trochanter is 
plesiomorphic-primitive in being less expanded than is typical of the well known late Cretaceous 
ornithomimosaurs. 

Ceratopsians are common late Cretaceous dinosaurs of North America and Asia with questionable 
records from Europe and South America. None have previously been reported from the early Cretaceous. 
An ulna recovered from early Aptian sediments of the Strzelecki Group at the Arch near Kilcunda, 
Victoria, Australia, shares the foreshortened and mediolaterally compressed form characteristic of 
ceratopsians and not known in other vertebrate groups. In particular, the Australian specimen is 
remarkably similar in both size and morphology to an ulna of Leptoceratops gracilis from the 
Maastrichtian of Alberta, Canada. 

The presence in the early Cretaceous of Australia of two dinosaur groups best known from the late 
Cretaceous of the Northern Hemisphere suggests that an hypothesis of either' s origin on the Gondwana 
continents should not be rejected out of hand. 

BARSBOLD, R. & OSMOLSKA, H., 1990. Ornithomimosauria. Pp. 225-248 in The Dinosauria' Ed. 
D. B. Weishampel, P. Dodson & H. Osmolska. University of California Press: Berkeley. 

The molar / premolar boundary in the macropodiforms: cracking the code of molar cusp 
patterns in the transitional zone 

RIDE,W. D. L. 1994 

Department of Geology, Australian National University, Canberra, Australian Captial Territory, 2600. 

The striking functional discontinuity at the molar/premolar boundary of many macropodiforms 
(kangaroos, wallabies and rat-kangaroos) is reflected in cusp pattern modification in the deciduous 
premolar and the first molar. Following recent analysis of this in Jackmahoneya, other propleopinines, 
and Hypsiprymnodon, the analysis is extended to the Macropodidae. 



228 ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 

The great Canowindra Devonian fish kill 

RITCHIE, A. 1994 

Australian Museum, Sydney, New South Wales, 2000. 

The richest fossil fish site in Australia, evidence of a unique Late Devonian mass fish-kill, was first 
discovered in 1956 near Canowindra, N. S. W. Using a 22 tonne excavator, Alex Ritchie relocated the 
fish horizon in January 1993. Preliminary excavations indicate that the site contains hundreds, probably 
thousands, of complete fish specimens. Five taxa have been recognised to date. Three types of armoured 
fishes, placoderms, are present. Antiarchs (Bothriolepis and Remigolepis) are dominant, forming about 
95% of the fauna. An arthrodire (Groenlandaspis) is present but is extremely rare. The fourth type 
present is a crossopterygian, an elongate, air-breathing, lobe-finned fish named Canowindra after fhe 
town near where it was discovered. The January '93 excavation turned up four new crossopterygian 
specimens of a new form, different from Canowindra. Two of these were giants, over 1 .5 m long. 

Alex Ritchie hopes to carry out a major excavation at Canowindra in June, using logistic support 
provided by the local shire council and enlisting local high school students as volunteers. The enormous 
scale and importance of this fossil site is now clear - there could be several hundred tonnes of rich fish- 
bearing slabs awaiting recovery, easily excavated and readily accessible in a roadside locality! The great 
Canowindra Devonian fish-kill provides a unique opportunity for a detailed population study on an 
entire fauna, killed suddenly and buried quickly. Given sufficient community support and funding 
Canowindra has enormous potential for a major tourist, educational and scientific attraction for central 
west New South Wales. 



Burnt ratite eggshell from Pleistocene aeolian sediments 

SMITH 1 , M. A., MILLER 2 , G. & VAN TETS U , G. F. 1994 

1. Department of Prehistory, Australian National University, Canberra, Australian Capital Territory, 
2600. 

2. Center for Geochronological Research, University of Colorado, Boulder, CO, 80309, United States of 
America. 

3. CSIRO Division of Wildlife & Ecology, Lyneham, Australian Capital Territory, 2602. 

The history of Genyornis newtoni (Dromornithidae: Aves) is complicated by uncertainty surrounding 
the date of extinction and role of human predation. Work in progress near Wood Point, on the eastern 
side of Spencer Gulf, S. A., has revealed a thin horizon of burnt Genyornis and Dromaius eggshell 
stratified within a Pleistocene dune core. Dates of 47± 5 ka have been obtained for the eggshell of both 
ratite species using the protein diagenesis (amino acid racemisation) method. These dates are amongst 
the most recent obtained for Genyornis in a large series of such determinations on eggshell across the 
southeastern sector of the arid and semi-arid zone and suggest that this species was extinct over a large 
part of its former range by ca. 40-50 ka. Work is in progress to verify the AAS results using other 
dating methods, including AMS C14 on the eggshell and TL on the dune sediments. Several features of 
the site suggest a human agency for the accumulation of the eggshell. 

Lake Callabonna: 'Veritable necropolis of gigantic extinct marsupials and birds' 

TEDFORD, R. H. 1994 

Department of Vertebrate Paleontology, American Museum of Natural History, New York, N.Y., 10024, 

United States of America. 

The famous fossiliferous deposit at Lake Callabonna, northern South Australia, lies at the base of the 
local Quaternary sequence in laminated clays and fine sands tentatively correlated with the Millyera 
Formation of nearby Lake Frome. At Lake Callabonna these deposits accumulated in a lake of variable 
salinity, several times the size of the present saltpan. Plant remains contained in the deposits indicate a 



ABSTRACTS: CAVEPS 1993. REC. S. AUST. MUS. 27(2) 229 

more arborescent flora than occurs near the lake today but one containing taxa still living in the region. 
This and stratigraphic evidence suggests a seasonal, yet wetter than present, climate with a fluctuating 
water-table. Geological correlations with Lake Frome suggest a medial Pleistocene age, 0.1-0.7 Ma, as 
the span during which the Callabonna fauna lived. 

The large-bodied vertebrates were mired while walking across the lake floor when the surface 
appeared dry but the underlying clays were water saturated. This mode of accumulation has yielded 
articulated skeletons of taxa known elsewhere only from fragments. In addition to the well-known 
remains of Diprotodon optatum and possibly a second, smaller species, there are three species of 
Sthenurus: a new giant form, the somewhat smaller S. tindalei, and the considerably smaller S. 
andersoni. Other kangaroos include: Macropus sp. cf. M. titan, a smaller Macropus sp., and 
Protemnodon sp. cf. P. brehus. Skeletons of Phascolonus gigas, the dromornithid Genyornis newtoni, 
and an emu similar to the living species are also present. 



The Lancefield megafauna site, Victoria 

VANHUET, S. 1994 

Department of Earth Science, Monash University, Clayton, Victoria, 3168 

The Lancefield Quaternary megafaunal site was the third megafaunal locality to be discovered in 
Australia. There are three separate sites located at Lancefield; the Classic, located in 1974, the South, 
located in 1983 and Maynes Site, first discovered in 1843 and subsequently relocated in 1990. The 
cause of the death of the megafauna at Lancefield is related to climatic changes occurring across 
Australia at the time of the collection. These changes had a detrimental effect on the habitats of 
browsing megafauna and their ecologically dependent carnivores and scavengers. Disease may have 
affected animals that were in a weakened state through lack of food and water. 

The presence of large numbers of bones in a concentrated area is due to transportational processes. 
The topographical nature of the area and the characteristic Quaternary climate encouraged major 
seasonal rainfall and sheet flooding. Bias in the collection is also due to transportational processes. 
Abrasion during transportation led to the destruction of the more fragile elements and weathering would 
have added to the bones fragility and subsequent fragmentation. Other biases in the collection, such as 
up to 90% of the elements being identified as the species Macropus titan, and the absence of juveniles 
of this species may well be related to the climatic conditions and subsequent habitat changes in the area 
at the time. 



Functional anatomy of the Lake Callabonna sthenurine kangaroos 

WELLS', R. T. & TEDFORD 2 , R. H. 1994 

1. School of Biological Sciences, Flinders University of South Australia, Bedford Park, South Australia, 
5042. 

2. Department of Vertebrate Paleontology, American Museum of Natural History, New York, N.Y., 
10024, United States of America. 

This paper reports on our functional anatomical study of three species of the extinct kangaroo 
Sthenurus from Lake Callabonna, northern South Australia. Complete skeletons of these animals were 
recovered from the lake sediments thereby providing an unparalledled opportunity for comparative 
anatomical study. The large taxa show sexual dimorphism. All differ from extant Macropus species in 
having short deep skull, long forefeet with reduced lateral digits, and functionally monodactyl hindfeet. 
The hand is modified for grasping, the forelimb may be raised above the head to reach high browse; the 
vertebral column shows limited flexion, but considerable extension in the anterior end; the pelvis is 
modified for flexion and adduction of the thigh; the hindlimb is more massive than Macropus species, 
and although of similar proportions there is greater emphasis on tendons and ligaments to augment 
muscular action. The mechanics of movement in these extinct species will be discussed. 



SOUTH 

AUSTRALIAN 

MUSEUM 



VOLUME 27 PART 2 
OCTOBER 1994 
ISSN 0376-2750 



PROCEEDINGS OF THE 4TH CONFERENCE ON AUSTRALASIAN 
VERTEBRATE EVOLUTION, PALAEONTOLOGY AND SYSTEMATICS 
(CAVEPS-93) ADELAIDE, 19-21 APRIL 1993 



CONTENTS: 



INTRODUCTION 
ARTICLES 



V J N. S. PLEDGE 

Fossils of the Lake: a history of Lake Callabonna excavations. 
79 R. H. TEDFORD 

Succession of Pliocene through medial Pleistocene mammal faunas of southeastern Australia. 
" J. LONG & B. MACKNESS 

Studies of the Late Cainozoic diprotodontid marsupials of Australia. 4. The Bacchus Marsh 

Diprotodons - geology, sedimentology and taphonomy. 
Ill J.A.MCNAMARA 

A new fossil wallaby (Marsupialia: Macropodidae) from the southeast of South Australia. 
117 N. S. PLEDGE 

Cetacean fossils from the Lower Oligocene of South Australia. 
125 T. H. WORTHY 

Late Quaternary changes in the moa fauna (Aves: Dinornithiformes) on the West Coast of the 

South Island, New Zealand. 
135 G. F. VANTETS 

An extinct new species of cormorant (Phalacrocoracidae: Aves) from a Western Australian peat swamp. 
139 W. E. BOLES & B. MACKNESS 

Birds from the Bluff Downs Local Fauna, Allingham Formation, Queensland. 
151 A.THULBORN 

Mimicry in ankylosaurid dinosaurs. 
159 A. W. WHITE & M. ARCHER 

Elseya lavarackorum, a new Pleistocene turtle from fluviatile deposits at Riversleigh, north-western 

Queensland. 
169 M. J. TYLER, H. GODTHELP & M. ARCHER 

Frogs from a Plio-Pleistocene site at Floraville Station, northwest Queensland. 
175 C. BURROW 

Form and function in scales of Ligulalepis toombsi Schultze, an enigmatic palaeoniscoid from the 

early Devonian of Australia. 
187 G. MCNAMARA 

Cape Hillsborough: an Eocene-OIigocene vertebrate fossil site from northeastern Queensland. 
197 B. MCGOWRAN & Q. LI 

The Miocene oscillation in Southern Australia. 

NOTES 
213 M.JINMAN 

World Heritage and fossils. 
215 P.CREASER 

Protection of Movable Cultural Heritage 

ABSTRACTS 

217 Other papers presented at CAVEPS-93, 19-21 April 1993. 

Published by the South Australian Museum, 
North Terrace, Adelaide, South Australia 5000.